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The Entomological Society of America 


VOLUME IV, 1911 



ITHACA, N. Y. Washington, D. C. 


GuELPH, Ontario, Canada. Boston, Mass. 


Boston, Mass. Philadelphia, Pa. 


Stanford Univ., Cal. Urbana, Ills. 

HERBERT OSBORN, Managing Editor, 
Columbus, Ohio. 




Notes on African Myrmeleonidae. Nathan Banks 1 

The Lithobiomorpha of the Southeastern States. Ralph V. Chamberlin. . . 32 
Notes on the Synonomy of the Genera Included in the Tribe Lachninae. 

H . F. Wilson 51 

Summary of the Food Habits of American Gall Midges. E. P. Felt 55 

The Structure and Systematic Importance of the Spermatophores of Crickets. 

J. P. Jensen 63 

Notes on the Life History of the Larch Case-Bearer (Coleophora Laricella). 

Glenn W. Herrick 68 

Further Biological Notes on the Colorado Potato Beetle Leptinotarsa 

10-lineata Say. A. A. Girault and James Zetek 71 

Minutes of the Minneapolis Meeting. J. G. Sanders 84 

Mayflies of Fall Creek. Anna H. Morgan 93 

Announcements of Further Results Secured in the Study of Muscoid Flies. 

C. H. Townsend 127 

Robberflies of the Genera Promachus and Proctacanthus. J. S. Mine 153 

Locomotion of the Larva of Calosoma Sycophanta. A. F. Burgess 178 

Notes on the Pear Slug. R. L. Webster. .' 181 

The Mechanism in the Hatching of the Walking Stick Diapheromera 

Femorata Say. H. H. P. and H. C. Severin 187 

Some Suggested Rules to Govern Entomological Publications. T. D. A. 

Cockerell 192 

The Composition of Taxonomic Papers. R. A. Muttkowski 194 

The Structure of the Central Nervous System of Corydalis Larva. 

W. A. Hilton 219 

A Structural Study of the Caterpillars II. The Sphingidae. W. T. M. Forbes 261 
Some Notes on Heredity in the Coccinellid Genus Adalia, Mulsant. Miriam 

A. Palmer 283 

Specific Characters Used in the Genus Pseudococcus. P. E. Smith 309 

Corrections to my paper in the June (1911) Issue of the Annals. C. H. 

Townsend 328 

Monograph of the Gall-Making Cynipidae (Cynipinae) of California. 

D. T. FULLAW.AY 331 

The Genera Hypera and Phytonomus in America North of Mexico. E.G.TiTUS 383 

VoJume IV. Number I. 



The Ejitomological Society of America 

MARCH, 1911 



ITHACA, N. Y. Washington, D. C. 


GuELPH, Ontario, Camaua. Boston, Mass. 


Boston, Mass. Philadelphia, Pa. 


Stanford Univ., Cal. Urbana, Ii,i,s. 

HERBERT OSBORN, Managing Editor, 
CoLtTMBUS, Ohio. 


Eatered as second clau matter April 1 1, 1908, at the Poit Office at Columbus, Ohio, 
under the Act of Congress o( March 3, 1879. 

The Entomological Society of America. 

TOXrNDMV 1906. 

ornoERs 1911. 

/>ej»<ie»t— Herbert Osborn Columbus, Ohio 

First Vice-President — Lawrence Bruner Lincoln, Nebraska 

Second Vice-President— h.T). MacGillivr.'W Ithaca, New York 

Secretary-Treasurer — A. D. MacGillivray Ithaca, New York 

Execulive Committee— -Tsu Officers, and J. H. Comstock, W. M. Wheeler, 
J. B. Smith, C. J. S. Bethune, Henry Skinner, A. D. Hopkins. 
Committee on Nomenclature— }i. T. Fernai.d, E. P. Felt, T. D. A. Cockerell. 

Price List of Publications. 

Annals, Vols. I, II and III, complete, each $3.00 

Annals, Vols. I and II, Parts 1, 2 and 4, each 1.00 

Annals, Vols. I and 11, Part 3, each 50 


Proceedings of first three meetings; Constitution, By-Laws and List of Mem- 
bers 25 

Wheeler, Wm. M. — Polymorphism of Ants 30 

Osborn, Herbert — The Habits of Insects as a Factor in Classification 20 

Severin, H. H. and Severin, H. C. — Aaatomical and Histological Studies 
of the Female Reproductive Organs of the American-Saw fly, Cimbex 
Americana, Leach >. 25 

Felt, E. P. — Some Problems in Nomenclature 10 

• Hammar, A. G. — On the Nervous System of the Larva of Corydalis comuta L. . .25 

Bradley, J. C. — A case of Gregarious Sleeping Habits among Aculeate 

Hymenoptera , 10 

Davis, J. J. — Notes on the Life History of the Leafy Dimorph of the Box- 
elder Aphid, Chaitophorus negundinis Thosi .10 

Hambleton, J. C.-^The Genus Corizus, with a Review of the North and 

Middle American Species 25 

Girault, a. A. — Biological Notes on Colorado Potato Beetle 25 

.^GiRAULT, A. A. — A Monographic Catalogue of the Mymarid Genus Alaptus. . . .25 

Severin, H. H. and Severin, H. C. — Internal Organs of Reproduction of 

Male Saw-fly 15 

Smith, C. P. — A Preliminary Study of the Axaneae Theraphosae of California. . .75' 

Davis, J. J. — Studies on Aphididae 20 

Riley, W. A. — Muscle Attachment of Insects 15 

Needham, J. C. — Critical Notes on the Classification of the Corduliinae 

(Odonata) 15 

Howard, L. O. — A Key to the Species of Prospaltella with Table of Hosts 

and Descriptions of Fovir New Species. 15 

Hood, J. D. — Two New Species of Idolothrips 10 


Biological Building, O. S. U., Columbus, Ohio. 



The Entomological Society of America 

Volume IV MARCH, 1911 Number I 


By Nathan Banks. 

The following article is based largely on the collection of 
African Myrmeleonidfe of the Berlin Museum, which the cura- 
tor, Dr. R. Heymons, wished me to work out for them. Mr. 
Esben Petersen has kindly loaned me his collection of Abyssin- 
ian Myrmeleonida?, which contains several species previ- 
ously unknown to me. My own collection contains a num- 
ber of species, principally from South Africa from Dr. H. 
Brauns' duplicates from the Petersen collection, and material 
purchased from dealers. 

There are many other species recorded from Africa. Some 
of these are probably synonyms of well-known species. I have 
made a new arrangement of the genera, and given tables to 
species as far as possible. 

Much of the Berlin Museum material is from German East 
Africa (D. O. Afrika), but many valuable specimens are from Ger- 
man Southwest Africa (D. S. W. Afrika), from Kamerun and 

The types of the new species, except where otherwise indi- 
cated are in the Berlin Museum. 


1. In the hind wings but one (rarely two) cross-veins before the origin of the 

radial sector; the anal usually ends in the margin (DendroleoninK).. . .11 

In the hind wings four or more cross-veins before the origin of the radial 

sector; the anal often runs into the cubital fork (Myrmeleoninoe) 2 

2. In the hind wings the anal is not connected directly to the hind margin by 

cross-veins, but to the post-anal by a series of cross veins; wings heavily 

marked; pronotum very much broader than long (Palparini) 3 

In hind-wings the anal vein is connected directly to the hind-margin by several 
cross-veins (Myrmeleonini) 6 

3. Two or more series of costal cells nearly to the base of wing Stenares 

But one series of costals, except near the stigma -t 

4. Apex of subcosta incrassate Pamexis 

Apex of subcosta not incrassate o 

2 Annals Entomological Society of America [Vol. IV, 

5. Antennae not their diameter apart at base; the basal joint with long bristles 

Antenna more than diameter of basal joint apart; smaller species. .Tomatares 

6. Spurs longer than the basal joint of tarsus, which is very short 7 

Spurs not longer than the basal joint of tarsus, which is longer than the 

second joint 9 

7. Body very hairy; legs very short; spurs much curved 8 

Body but little hairy; legs more slender; spurs nearly straight; a line through 

the apical part of wings Myrmecaelurus 

S. Costal series double Syngenes 

Costal series single Acanthaclisis 

9. The branches of radial sector are bent to form a straight line through the 
middle of the apical part of the wing; basal joint of the tarsus about as 

long as the apical Nesoleon 

No such line through the wings 10 

10. In fore wings the radial sector arises before the cubital fork; many eostals 

before the stigma are crossed Hagenomyia 

In fore-wings the radial sector arises much beyond the cubital fork; eostals 

are mostly simple Myrmeleon 

(a) A series of cross-veins before the radial sector are crossed; very large 

species subgenus Macroleon 

11. Spurs present and distinct 12 

No .spurs: 

Legs very slender Gymnocnemia 

Legs of usual length Gymnoleon 

12. The forks of cubitus in the fore-wing are parallel for a distance, and the anal 

is also parallel to them Creagris 

The forks of cubitus diverge, and the anal is not parallel to the upper branch. 
and to the lower only for a short distance 13 

13. Joints 2, 3, and 4 of tarsus very long, each as long as basal joint. , . .Megistopus 
Joints 2 and 3 very short, but little longer than broad 14 

14. Legs very slender; the tibiEe I and II as long or longer than femora I and II; 

basal tarsal joint nearly as long as apical; spurs as long as two joints; 

antennsE very slender; w'ngs broad and much marked (Dendroleoni)... .15 
Lefis shorter; the tibi.p I and II shorter ihan femora I and II, the basal tarsal 

joint plainly shorter than the apical (Nemo'eonini) Ill 

l.T. Wings especially hind pair very stronj^ly ;alcate at tip; first joint of tarsus 

longer than the last Cymothales 

Wings hardly falcate; the branches of radial sector bent to form a straight 

line through the middle of apical half of wings Dendroleon 

16. In hind-wings the anal does not run directly to the margin but bends upward 

making a long curve; the radial sector arises before cubital fork in buth 

wings Echthromyrmex 

In hind-wing the anal runs directly to the hind margin 17 

17. Spurs but little longer than the long first joint of tarsus; first branch of radial 

sector arises beyond end of the anal vein Nemoleon 

Spurs longer than the first tarsal joint, which is short IS 

18. Spurs as long as three or four joints of tarsus Formicaleon 

Spurs not so long, about as long as two tarsal joints Macronemurus 

Under Stenares I have placed Crambomorphus. Both are 
hardly more than subgenera of Palpares. Symmathetes is 
united with Palpares. A species of Glenurus is described from 
Africa, but from the figure it is a Dendroleon; the genus Palpari- 
diiis, lately described by Peringuey, appears to be identical with 
Echthromyrme.x. The Centroclisis of Navas I consider the same 
as Acanthaclisis. 

1911] Notes on African MyrmeleonidcB 3 

Palpares Rambur. 

Of this genus there are about forty named species from 
Africa; doubtless there are several more. They are the grandest 
and most beautiful insects of the family, if not of the entire 
Neuropteroid series. Their marking are however, variable, 
as may be easily seen from even a small series of specimens. 

I have tabulated thirty-four of the species, but a few are 
perhaps synonyms. 

Type — P. libelluloides. 


1. Hind-wings black, except seven or eight small spots 2 

Hind-wings much more pale, only with dark bands 3 

2. Hind-wings very narrow, lanceolate karrooanus 

Hind- wings broad, not lanceolate voeltzkowi 

3. Fore-wings with a complete band across before middle 4 

Fore-wings without complete band before middle, outer margin of wings 

scarcely sinuate 6 

4. Outer margin of wings sinuate; very large species (Symmathetes) .5 

Outer margin of wings not sinuate, rather small species ovampoanus 

5. Basal band of fore-wings reaches hind margin, apical spot divided; hind- 

wings with the stigmal spot furcate in front, apical spot divided.. . .gigas 

Basal band of fore-wings does not reach hind margin, apical spot not divided; 

hind-wings with the stigmal spot not furcate in front, apical spot not 

divided, but emarginate moestus 

G. A large dark spot over the cubitus in the hind wings 21 

Not more than a dot over the forking of cubitus, although a dark spot may be 
near by 7 

7. Hind wings with complete bands 8 

Hind wings without complete bands 13 

8. Fore-wings densely reticulate with black, but leaving three clear yellow bands. 

flavofaciatus and genialis 
Fore-wings without three clear yellow bands 9 

9. Median band with a projection toward the cubital fork, or else a separated 

spot near by, the stigmal band with an upward projection as well as one 

behind 10 

Not so marked li 

10. Median band of hind-wings connected to the stigmal, and median has a pro- 

jection toward cubital fork latipennis 

Median band well separated from stigmal, and a spot near cubital fork sepa- 
rate from the median band inclemens 

11. Median and stigmal bands of hind wings well separated insularis 

Median and stigmal bands connected or nearly so 12 

12. Apical spots of both wings divided so as to form an anterior and posterior 

apical streaks elegantulus 

Apical spots not so divided amitinus 

13. Wings with all small spots, no large ones; legs yellow sparsus 

Wings with some large spots 14 

14. Outer margin of both wings very narrowly black; no marks between median 

and stigmal spots in fore-wings 15 

Outer margin of wings with dots or spots; some marks between median and 
stigmal spots in the fore-wings 16 

15. A black band below antennae segrotus (tessellatus) 

No black band below antennae tigris 

16. Median spot of hind-wings not reaching one-half way across wing; in fore- 

wings the median and stigmal spots are small 17 

Median spot of hind-wings reaches one-half way across wing 18 

4 Annals Entomological Society of America [Vol. IV, 

17. Legs yellowish; few small spots in fore-wings; (female) submaculatus 

Legs blackish, more small spots in fore-wings, (male, probably same as 

submaculatus) nyicanus 

IS. In fore-wings the median spot is barely larger than the numerous other spots. 

In fore-wings the median spot is much larger than the many small spots. . . .19 

19. Many small spots in the basal part of the hind-wings furfuraceus 

Few, if any, small spots in the basal part of the hind wings 20 

20. Larger; spots not broken up much tristis 

Smaller, wings shorter, and appendages shorter, stigmal spot of hind-wings 

more broken up Interioris 

21. Basal spot of hind-wings reaches nearly to base of wings 22 

Basal spot of hind-wings not reaching toward base 23 

22. Basal spot large and angulate, also large stigmal spot immensus 

Basal spot in form of a streak, also other streaks radiatus 

23. Median band of fore-wings very small, reticulate, with darker edges, fore- 

wings often yellowish 24 

Median band of fore-wings distinct, reaching one-half way across wing. . . .27 

24. Median and stigmal spots of hind-wings connected; but few small spots in 

the fore-wings stuhlmanni 

Median and stigmal spots not connected, more small spots in the fore- wings. 


2.5. Hind part of fore-wings not tessellate with dark spots dubiosus 

Hind part of fore-wing tessellate with dark spots 20 

26. Fore-wings not very yellowish speciosus 

Fore-wings plainly j'ellowish cafEer 

27. Median band of hind-wings bent inward behind; stigmal spot broader behind 

than in front, and reaches hind margin 28 

Median band not bent inward at tip; stigmal spot not reaching hind margin 
and narrow behind 32 

28. Wings broad; median band of hind-wings does not reach the hind margin. 

Wings usually narrow, median band of hind-wings reaches hind-margin. . . .29 

29. Apical mark of hind wings connected to the stigmal 30 

Apical mark of hind-wings not connected to stigmal; two yellowish spots on 

front of the pronotum 31 

30. But lightly marked, stigmal spot of fore-wings larger, from Madagascar. , 

More heavily marked, stigmal spot smaller; from South Africa (may l)c same) 


31. No small spots between median and stigmal spots in fore-wings. . . .damarensis 
Many small spots between median and stigmal spots in the fore- wings. . . . 


32. Stigmal spot of hind-wings connected to a large spot just before it . . . cephalotes 
Stigmal spot of hind-wings without a spot just before it 33 

33. Stigmal spot of hind wings reaches across wing, in fore-wings nearly across; 

median spot of fore- wings very narrow percheroni 

Stigmal spot of hind- wings and fore-wings not near across wing. . .libelluloides 

Palpares formosus n. sp. 

Head black, clypeus yellow; pronotum black, with two yellow spots 
in front; rest of the thorax black, with several pale spots through the 
middle; legs black; abdomen pale, darker toward tip. Wings very 
slender; forewings densely spotted with small brown dots, but a narrow 
angulate median band crosses the wing, the stigmal spots small, and a 
rather large preapical spot. Hindwings with a large spot over the fork 
of the cubitus, an angulate median band crosses the wing to the hind 
margin, leaving two small hyaline spots on the hind margin; a stigmal 
band crosses the wing, leaving one hyaline spot on the hind margin, 

1911] Notes on African Myrmcleonidcc 5 

and it has an inner projection toward the median band; a transverse, 
preapical band, connected each side around the tip and thus enclosing 
a hyaHne spot. 

Expanse SO mm. 

From Willowmore, Cape Colony, 20 Febr. (Dr. Brauns). 
Similar to P. damarensis McLach., but with more slender wings; 
the forepair with spots all over the surface, not clear between 
the median and stigmal bands. (Banks coll.) 

Acanthaclisis Rambur. 

This genus is related to Myrmeleon, but distinguished by the 
very heavy legs, and the much curved or rather geniculate 
spurs; the body and legs are extremely hairy; the antennae are 
rather close together at base ; the wings in most forms have a line 
through the apical part of the wings formed by the bending of 
the branches of the radial sector, and there is a similar line 
through the cubital area formed by the bending of the branches 
of the upper cubitus. 

Type — A . occitanica. 

The genus Syngenes formed for A. debilis Gerst, is scarcely 
more than a subgenus; it has many of the costal crossveins 
crossed.* The genus Centroclisis is, I think, a synonym of 
Acanthaclisis, its type species agreeing closely with my speci- 
mens of A. distificta. 

Quite a nuriiber of species have been described from Africa; 
Dr. Van der Weele examined Gerstaecker's types and placed 
most of them as synonyms of other forms; I have seen only a 
few of the species, and the following table is based partly on 
the descriptions. 

1. Costal area with many of cross veins crossed (Syngenes) longicornis 

Costal area with most of cross-veins simple or forked 2 

2. A black streak through middle of fore-wings lineatipennls 

No black streak in fore-wings 3 

.3. Spurs evenly curved dasymalla 

Spurs geniculate -1 

4. Fork of cubitus with a distinct spot mashunensis 

Fork of cubitus immarked .5 

.5. Various patches of white cross-veins in both wings; many small dark dots; 

larger ones along upper edge of cubitus, especially at end gulo 

Few patches of white cross- veins; less marked with dark, wings rather more 

acute at tips 6 

6. — Costal series almost all simple; pronotum not so plainly lineate with black. , 

Costal series mostly forked; pronotum very distinctly lineate with black. . 


*A. americana, A. fallax have many costals forked; .-I. japonica, A. horrida, 
and some Australian species have many costals crossed. 

6 Annals Entomological Society of America [Vol. IV, 

Acanthaclisis baetica Rambur. 

Not before recorded from Africa; in the Petersen coll. a 
specimen from Oran, Algeria agrees with Spanish specimens. 

Myrmecaelurus Costa. 

This genus is related most closely to Nesoleon, but separated 
from that as well as from Myrmeleon by the longer spurs; there 
is a line through the apical part of the wing. 

There are several species recorded from Africa, but I have 
seen but one which is new. The Myrmeleon tristis has been 
referred to this genus, but is nearer to Myrmeleon, and is better 
placed in a new genus. 

Type — M. flavus { = trigrammus). 

Myrmecaelurus subcostatus n. sp. 

Pale yellow; no mark between antennae, and only very faintly on 
each side on the anterior part of vertex; antenna pale brownish; pro- 
notum with a black line each side not reaching cither margin, and a 
median black dot on front margin; three black lines on rest of thorax, 
the median one nearly complete (single on the scutelli), the lateral ones 
interrupted, some dark lines over base of coxae I and 11; abdomen with 
a dark median stripe from base to tip, and a dark stripe each- side on 
the venter. Wings hyaline, venation yellowish, the subcosta very 
strongly marked with black nearly to the stigma; the black extending 
up on each costal crossvein; the radial sector much marked with black, 
and at tip with a black streak extending out over the end of radius 
toward the tip of wing; some dark dots on origin of branches of cubitus, 
and a few others near middle of wing. In the hind-wings the subcosta 
and radius are marked near base, and to a lesser degree on the radial 
sector and cubitus, and a few veins near middle of wing. Pronotum 
much broader than long, rounded in front. Wings rather broad near 
tip; about seven or eight crossveins before radial sector, and about ten 
branches to the radial sector; costals simple; hind-wings a little narrower 
but scarcely more acute than the fore-wings. 

Expanse 58 mm. 

From Erythraea (Kristensen coll.); types in coll. Banks and 

Nesoleon Banks. 

This genus is near Myrmeleon, having several crossveins 
before origin of the radial fork in hind wings, but a single costal 
series, and the spurs no longer than basal tarsal joint. There 
are from 6 to 10 crossveins before radial sector (more than in 
Myrmeleon, and the spurs or one of them is much shorter than 
the basal tarsal joint, which is nearly as long as the apical joint; 
the antennae are wide apart at base. The wings are broad and 

1911] Notes on African Myrmeleonidce 7 

subfalcate at tip, and differ at once from Myrmeleon in having 
a line through the apical part of each wing, partly formed by 
the bent branches of the radial sector. 
Type — N. braimsi Bks. 

1. Wings with large black spots 2 

Wings with only small black marks 3 

2. Wings almost all black, more than apical half of hind-wings black, only the 

stigma and an apical spot pale braunsi 

Wings mostly pale, less than apical half of hind-wings black, large stigmal 
and apical spots pale boschimanus 

3. No interantennal mark 4 

Interantennal mark distinct 5 

4. Pronotum with three dark stripes (unnamed species from Kamerun, but one 

specimen) sp. 

5. Pronotal stripes slender and broken; wings rather narrow; a dark spot at the 

base of the stigma, one at end of the cubital fork, and one over cubitus 

and median near their ends; elsewhere but little marked trivirgatus 

Pronotal stripes plain and complete 6 

6. Wings rather evenly marked all over with small blackish spots at the forks 

of the veins; the outer gradates not more prominent than other veins, nor 

a mark at the end of the cubital fork punctatissimus 

Wings not evenly marked all over 7 

7. Wings almost all hyaline, only faintly marked and then most noticeable near 

the outer apical margin; stigma darker at base pallens 

Wings unevenly marked, the stigma with a prominent dark spot at base, the 
outer gradates marked, and a pale unmarked streak beyond it, no promi- 
nent spot at end of cubital fork mysteriosus 

Similar to N. mysteriosus, but the wings not much marked, except at the 
gradate series, and near the end of the anal vein, thus two oblique lines 
on each fore- wing; stigma dark at base; longitudinal veins of wings strong!)' 
marked variegatus 

Nesoleon braunsi Banks. 

I have seen only the types from Willowmore, Cape Colony, 
(Dr. Brauns). 

Nesoleon boschimanus (Peringuey). 

Head black; a large pale spot each side on face; vertex pale, with 
black dots, three in each of three rows, median spots sometitnes con- 
nected; pronotum with broad median black, and two narrower stripes 
each side, the median continued back over the thorax, and the lateral 
also continued, but in a broken and branched form; abdomen black, 
some of the segments beyond middle are pale at tip; femora brown; 
tibicB pale, with a black tip, and on front and middle pair a median band 
also; tarsi, except basal joint, mostly black; spurs weak, hardly more 
than one-half the length of the basal tarsal joint. Wings moderately 
broad, about as in N. mysteriosus; apex acute; venation interrupted 
black and white; forewings with larger spots along hind and outer 
margin, a sub-basal spot, a larger oblique median spot, a larger stigmal 
spot, sloping inward, a spot behind this one; hind wings pale on basal 
three-fifths; two small spots near disc of wing, then a broad prestigmal 
band, narrowed in the middle, and broadened behind and reaching out- 
ward to near tip of wing, a prominent spot beyond the stigma, nearly 

8 Annals Entomological Society of America [Vol. IV, 

connected to the large black spot behind it. Seven cross-veins before 
radial sector in hind wings; eight branches to radial sector in both 
pairs; abdomen shorter than wings. 
Expanse 40-4,5 mm. 

From Deutsch Sud. W. Afrika; Grootfontein, Okahandya, 
and Gr. Namalaut. 

Nesoleon mysteriosus (Gerst.). 

From various places in D. O. Afrika (Berlin Museum); 
and Mt. Algego, Uganda (Banks coll.) 

Nesoleon variegatus (Klug). 

From Harrar, Abyssinia (Petersen and Banks coll.) 

Nesoleon trivirgatus (Gerst.) 

From Grootfontein, D. S. W. Afrika. This may be but a 
pale form of N. variegatus. 

Nesoleon punctatissimus (Gerst.) 

From Windhoek, Reitfontein and Reheboth, D. S. W., 
Afrika, and Lindi, D. O. Afrika. This may be the MyrmeJeon 
lanceolatus Rambr. 

Nesoleon pallens. (Klug). 

Several specimens collected by Kristensen in Erythraea 
I believe are this species; there is some variation in length and 
breadth of wing and one specimen has the wings about as Klug 
figures, but the abdomen does not show the transverse dark 
marks, nor does Klug's description mention them, so I presume 
the figure is too highly colored in this respect. Klug's species 
is surely a Nesoleon as he shows plainly in his figure the fine line 
in the apical venation of the forewings. 

Hagenomyia new genus. 

This genus agrees in general with Myrnieleon ; there are sev- 
eral crossveins before the radial sector in the hind w4ngs; the 
spurs are no longer than the first tarsal joint; which is longer 
than the second; the antennae are wide apart at base; and there 
is no line through the apical part of the wings; it differs from 
Myrmeleon in the much broader wings, in having the radial 
sector of forewings arising before the cubital fork, and in 
having many costals before the stigma crossed. 

Type — Myrmeleon tristis Hagen. 

1911] Notes oil African Myrmeleonidce 9- 

Hagenomyia tristis (Haj,'cn). 

This is very common in many parts of Africa; I have seen 
specimens from Argabba, Tewe, N. Usambara, Kamerun, 
Togo, Kongo, Tanganjika, Sansibar, Angola, Madagascar, 
and various places in D. O. Afrika (Berhn Aluseum), Erythraea, 
Abyssinia, and Abutshi, Niger (Banks coll.) 

Myrmeleon Linnaeus. 

In this genus there are several crossveins before the radial 
sector in both wings, the anal is connected to margin; there is 
no straight line through the apical part of the wings, the basal 
joint of tarsus is much shorter than the apical joint and the spurs 
are not or but a little longer than the basal joint ; the costals are 
simple, the cubital forks diverge, and the wings are usually narrow. 

Type. — Myrmeleon formicarium Linn. 

There are two subgenera; Myrmeleon and Macroleon, the 
latter with large species, mostly broader wings, and with several 
of the crossveins before the radial sector crossed ; the type of this 
subgenus is M. (Myrmeleon) validus McLach. 

Many species have been described from Africa, several of 
which probably do not belong to the genus as here restricted, 
and others are perhaps synonyms of some of the common 
species. Those that I have seen are tabulated below. 

1. Fore-wings with an oblique dark mark up from the end of the anal vein, and 

dark spots on the cubitus and median near the tip; pronotum dark. . . . 

No such marks 2 

2. Outer apical margin of both wings narrowly dark; pronotum dark 

Outer apical margins not dark '. 3 

3. A large dark cloud before and beyond the large white stigma lynceus 

No such cloud 4 

4. Abdomen plainly banded with pale; pronotum dark, indistinctly marked. . (i 
Abdomen not plainly banded with pale; pronotum pale yellowish, at least on 

sides h 

5. A simple, narrow median dark stripe on the pronotum medialis 

A simple median, and a lateral stripe for most of the distance on pronotum. , 


A broad median stripe furcate in front on the pronotum furcatus 

A broad median dark stripe on pronotum, serrate on the sides and occupying 
most of the surface lethifer 

6. Size small, stigma barely distinct; no series of crossed veins before radial 

sector in fore-wings, cubitus marked with black and white obscurus 

Size very large, stigm.i plainly white; a series of crossed veins before radial 
sector in fore-wings (Macroleon) V 

7. Wing tinged throughout with a vinous color validus 

Wings not tinged with vinous 8 

8. Interantennal mark includes a pale spot below antennse; black on clypeus; 

pronotum with five pale spots qulnquemaculatus 

Interantennal mark not complete below, but with a deep median indentation 
of pale; hardly black on clypeus; larger and with narrower wings atlas 

10 Annals Entomological Society of America [Vol. IV, 

Myrmeleon obscurus Rambur. 

Widely distributed in tropical and South Africa. Specimens 
have been examined from Harrar, Abyssinia (Petersen coll.); 
Tamatave and Vohemar, Madagascar, (Banks coll.) and 
Erythrsea; from Bagamoyo and Lindi, D. O. Afrika, Windhoek, 
D. S. W. Afrika, Livingstone, S. Afrika, and Kamerum (Berlin 

Myrmeleon lethifer Walker. 

From White River, Transvaal, Kibwezi, Brit. O. Afrika, 
and Kwidjwi, Ost Afrika. M. nigridorsis Kolbe is the same 

Myrmeleon doralice n. sp. 

Face yellow, a large shining black interantennal mark, curving 
e\'enly downward on the face, and abo\'e reaching to middle of vertex, 
with a median extension on vertex, and a black streak each side; anten- 
nae rather long, blackish, tip paler, pronotum longer than broad, nar- 
rowed in front, pale yellow, a median black stripe, widest in front, and 
a curved black streak each side from behind the transverse furrow; 
rest of thorax with middle black spot, and oblong spots over the base 
of wings, a basal median black spot on the scutelli; pleura black through 
the middle; abdomen shorter than wings, black, tips of segments nar- 
rowly pale, sides pale; legs pale yellowish, tips of tibiae rather darker, tip 
of last tarsal joint black; spurs not as long as basal tarsal joint, which 
is much shorter than the apical joint. Wings hyaline; venation pale 
yellowish, subcosta, radius, and cubitus marked with dark, stigma 
hardly distinct. Wings rather slender, acute at tips; about 10 cross- 
veins in forewings before radial sector, 6 in hind wings, 10 branches of 
radial sector; in the forewings the first fork of radial sector is so con- 
nected up to the radius as to appear as a distinct sector of the radius, 
thus apparently two radial sectors. 

Expanse GO mm. 

From Windhoek, D. S. W. Afrika. 

M3rrmeleon alcestris n. sp. 

Head mostly black, margin of face yellow, a median yellow spot on 
the clypeus, two dots between the antennae, and the orbits yellowish; a 
pale stripe across vertex in front, a large yellow spot each side, and three 
longitudinal submedian lines on the vertex pale, a spot each side on occiput 
pale; basal joints of antennae pale, with dark rings, rest dark brown; 
pronotum short, a median dark streak, a spot near each anterior comer, 
and a larger spot in each hind comer, dark; rest of thorax dark, with 
indistinct black marks, and the hind margins of the segments pale; 
coxae black, femora black, except pale base, tibiae pale, heavily spotted 
or streaked with dark; tarsi pale brownish, tip of last joint black; spurs 
not as long as basal tarsal joint, which is about two thirds as long as the 

1911] Notes on African Myrmeleonidce 11 

apical joint. Wings hyaline; venation black and white, the crossveins 
with black dots, the longitudinal veins with longer black streaks, a 
black spot at the base of the stigma, an oblique blackish cloud up from 
the end of the anal vein, and a spot over cubitus and median at the 
usual point for such spot. Wings slender, acute at tips; 7 cross-veins 
before radial sector in forewings, 5 in hind wings; 10 branches of radial 
sector in both pairs. 
Expanse 54 inin. 

From Reitfontein, D. vS. W. Afrika. 

Myrmeleon medialis n. sp. 

Head with a large black interantennal mark reaching down over 
most of the face and up to the ^'ertex, a dark mark on clypeus, in middle 
connected upward to the interantennal mark; vertex with a double 
median spot in front, a small median one behind, and a long one each 
side, black; antennas dark brown, short; pronottmi but little longer than 
wide, pale, with a broad median brown stripe; thorax pale, with some 
large median black spots, and one over base of each wing, jjleura with 
a broad broken black streak; abdomen black, with short white hairs, 
not as long as wings; legs pale, rather more brownish toward tips, with 
black bristles, tarsi quite long, first joint hardly as long as second and 
third together, fifth one and a half times as long as the first, spurs not 
as long as the first joint. Wings hyaline, venation pale yellowish, 
subcosta slightly marked with black, stigma barely distinct. Wings 
long and slender, forewings sub-falcate at tip, about 9 crossveins before 
radial sector in forewings, and (i in hindwings; 10 branches of radial 
sector in both wings. 

Expanse 60-68 mm. 

From Natal and Georgetown, Cape Colony. 

Myrmeleon furcatus n. sp. 

Head pale yellow, a black mark at base of each antennae, and a 
median vertical line; a broad, dark band above antennae; vertex with a 
pair of broken submedian stripes, a broader short stripe each side 
behind, with lateral projections toward eyes, and a lateral spot in front; 
antennse with a brown mark on basal joint; a narrow brown ring 
on second joint, beyond that brown; pronotum pale, with a broad med- 
ian dark stripe, furcate in front; a short dark stripe on each hind comer; 
rest of thorax dark, with hind margin of meso- and meta-thorax pale, 
a narrow pale line on each lateral lobe of the mesothorax, a spot on each 
lateral lobe of the metathorax, and the scutelli with a pale dot each side, 
and sometimes other spots; abdomen dark, pale at tip; legs pale, a dark 
line on the tibi£e, and a streak near tip of femora; spurs as long as the 
first tarsal joint, which is not as long as the next three together, and 
shorter than the last joint. Wings hyaline; venation pale, subcosta, 
radius and cubitus interrupted with dark, also slightly on the radial sec- 
tor, stigma dark, with a darker spot at base. Wings long, acute at 

12 Annals Entomological Society of America [Vol. IV, 

tips, the hind pair ahiiost subfalcate at tig; about eleven crossveins 
before the radial sector in forewings, six in'hintf wings; about twelve 
branches to radial sector in each wing. 
Expanse 73 to 90 mm. 

From Erythraea (Kristensen coll.). Types in collections 
of Banks and Petersen. 

Myrmeleon lynceus Fabr. 

From Victoria, Kamerun, and a smaller specimen from Togo. 
The Af. leucostigmatiis Weele, from Kamerun, is a synonym. 

Myrmeleon formicaroides Weele. 

One specimen from Misahohe, Togo. 
Myrmeleon quinquemaculatus Hagen. 

A common African species, specimens from Ginda (Petersen 
coll.), Kilimandjaro (Banks coll.) and various places in Togo 
and D. O. Afrika. M. polyzones Gerst. and M. rapax Kolbe . 
are synonyms as already stated by A^an der Weele. 
Myrmeleon validus McLachlan. 

From Tamatave, Majlagascar (Banks coll.). 

Myrmeleon atlas n. sp. 

Similar to M. o-maculatus, but much larger. Face yellow, with a 
median black spot below, the interantcnnal mark is not complete below, 
but indented with the pale reaching up between antennae, and the mark 
is not continued all around the lower edge of the antennal socket: 
marks of vertex similar to those of M. 5-maculatus; the pronotum 
shows a large pale median spot in front, and at each upper corner, with 
a narrow extension baclavard; the abdominal segments are pale only 
near middle (not to base as in .5-maculatus); the legs yellowish brown, 
not plainly marked. Wings much more slender than in M. .5-maculatus, 
especially the hind pair, and more falcate at tip; venation mostly dark, 
subcosta and radius plainly interrupted with pale; stigma not very 
distinct; there is a longer series of crossed veinlets before the radial 
sector in forewings than in M. .5-maculatus. 

Expanse 12G mm. 

From Kwidjwi, Ost Afrika. 

Gymnoleon new genus. 

But one crossvein before radial sector in the hindwings,. 
several in forewings ; costal series simple, cubital forks not very 
much divergent; antennae rather close together at base; pro- 
notum slender; no spurs; legs of moderate length, the last joint 
of tarsus as long as the basal joint. 

Type — G. exilis n. sp. 

By absence of spurs it is related to the genus Cymnocnemia, 
but differs at once in the very much shorter and stouter legs. 

1911] Notes oil African Myrmeleonidce 13 

Gymnoleon exilis n. sp. 

Face pale yellow, a large black interantennal mark, a row of spots 
across the vertex, the middle one the largest; pronotum pale yellowish, 
a pair of black submedian stripes, not quite parallel, and a lateral stripe 
each side; rest of thorax pale, with a pair of dark submedian lines, 
broken, but extending over the scutelli, a lateral dark line over base of 
wings; abdomen rather pale, with dark stripe on each upper side; legs 
pale, middle and apical mark on tibia, except the hind tibiae have only 
the apical mark, tips of tarsal joints black. Wings hyaline, venation 
mostly brown, but the longitudinal veins are interrupted with pale, 
many of the cross-veins with a pale dot or space; many of the forks in 
posterior and outer part of fore-wings have a black dot, a larger spot 
near end of anal vein, and faintly dark at anastomosis of cubitus and 
median, this last mark is more prominent in the hind-wings, often 
elongate. Wings rather slender, acute at tips, the hind pair plainly 
longer than the forewings, narrower and subfalcate at tips; about seven 
croSsveins before radial sector in forewings, about S branches of radial 
sector in each wing, in forewings the anal vein ends before the origin of 
first branch of radial sector, and the forks of cubitus are plainly diver- 
gent. The pronotum is longer than broad, and broadest behind; the 
abdomen is much shorter than the wings. 

Expanse 45 mm. 

From Ari Ugri, Erythrasa (Kristensen coll.) and from Lindi, 
D. O. Africa (Berlin Mus.). Type in coll. Banks; cotypes in 
coll. Petersen and in Berlin Museum. 

Gymnoleon elizabethae n. sp. 

Face yellow, a black band Ijelow the antennae reaching above them, 
the vertex broadly rounded in front and dark bro\\'n, back of vertex 
gray, with two rows of blackish dots, front of vertex with a median furrow 
making the front plainly bilobed ; antennae brown, tip paler, pronotimi 
short, dull blackish, a faint paler spot each side in front, and one or two 
in the middle; thorax brown, indistinctly marked with pale; a deep 
black band on metanotuni; abdomen blackish; legs short, yellow-brown, 
with black marks on tibiae, hind tibia only at tip, tarsi mostly dark 
brown, except the basal joint of hind tarsus which is yellow; no spurs; 
basal joint of tarsus as long as second and third together. Wings with 
black and white venation, on radius the black and white are in long 
streaks, elsewhere in short spots, along cubitus and its fork are several 
small clouds of brown, and a longer oblique mark on cubitus and median 
near their tip, stigma dark at base, and a cloud near end of anal and 
cubital fork. Wings slender, subfalcate at tips, anal and cubital fork 
in forewings run parallel for a distance longer than width of wing, the 
cubital fork nearly parallel to the cubitus; twelve branches of radial 
sector in each wing; in apical part of wings the longitudinal veins are 
very close together. 

Expanse 52 mm. 

From Port Elizabeth, Cape Colony, 25 Jan., (Brauns), 
(coll. Banks). 

14 Annals Entomological Society of America [Vol. IV, 

Creagris Hagen. 

This genus is known by the parallel forks of the cubitus in 
the anterior wings; there are but two rows of cells between the 
forks and these are separated by a nearly straight vein ; the anal 
vein also runs parallel to the cubital forks for some distance. 

Type — C. plumbeiis Oliv. 

Many species are known by the names, but several of these 
names apply to variations of C. plumbeus. Whether these 
forms should be kept distinct or all united under the one name 
must be determined by larger collections from representative 
parts of Africa. 

The forms I have examined are distinguished as follows: 

1. Venation pale, unmarked 2 

Venation marked with black and white 5 

2. Apex of hind-wings with a fuscous streak .3 

Apex of hind wings urunarked 4 

3. Very large species diana 

Species of moderate size nubifer 

4. Pronotum with indications of two narrow submedian lines; femora dotted 

with black murinus 

Pronotum with a median stripe or absent; femora scarcely dotted with black. 


5. Wings with black longitudinal streaks nlgrostrigatus 

Wings without streaks G 

6. Four large black spots along the radial sector in the fore-wings. . . . pretlosa 
No such large spots 7 

7. No black band below antennas S 

A black band below antennae, vertex with a frontal ridge, venation strongly 

marked with black and white aegyptiacus 

8. An oblique dark mark at end of anal vein in fore-wings; venation very plainly 

marked with black and white mortifer 

No marks on wings; venation only faintly marked with dark and pale. . . 


Creagris pretlosa n. sp. 

Face pale, a median vertical black line, and a brown spot tinder 
each antenna; antennas brown, base yellowish; vertex with a black band 
in front, a row of connected black dots, and behind are six separated 
black dots, the submedian ones being longitudinal; pronotum gray, 
black on the sides and a submedian pair of black stripes, broadest in 
front ; thorax gray, black marks on the lateral lobes and the usual black 
band on the mesonotum; abdomen black above, pale beneath. Legs 
brown, with much gray hair, femur I black on inner tip, tibia I black 
in middle and near tip, middle leg similar, hind legs paler, the tibiae 
black only at tij), all tarsi black, except yellow on basal and most of 
apical joint of tarsus I. Wings hyaline, venation black, interrupted 
with pale, the cross- veins mostly wholly black; four large black spots 
along the radial sector of forewings, another smaller at base of the 
stigma, a black streak along base of the cubital vein, small black spots 
at forking of cubitus and farther out on cubitus, one on a crossvein 
connecting median to the first branch of the radial sector, and one at 

1911] Notes on African MyrmeleonidcB 15 

end of the anal vein, the outer gradate veins fonn a black Hne parallel 
to the outer margin ; hindwings with a black spot at the base of the stigma 
one on a crossvein below and a little before it, and two faint ones on 
other crossveins from radial sector to radius, another near the end of 
the cubitus. Forewings broad, acute at tip, almost falcate, the costal 
margin at base plainly concave, the anal runs parallel to the fork of 
cubitus for much less than the width of the wings. Hind wings narrow, 
subfalcate at tip; eleven branches of radial sector in both pairs. 
Expanse 78 mm. 

From Detttsch Ost Afrika, Bex Lindi. (Berlin Museum). 

Creagris diana Kolbc. 

Of this magnificent species there are specirriens from Lindi, 
and Mohorovi, D. O. Afrika. Peringuey's recent species, 
C. incUtiis, seems to be the same form; his description agrees 
• with the East African specimens. 

Creagris nubifer Kolbe. 

Specimens from Lindi, D. O. Afrika; Bismarcksburg, Togo, 
and from Harrar, Abyssinia (Petersen coll.). Peringuey's 
recent C. proximus from South Africa agrees with these speci- 

Creagris plumbeus (Oliv.) 

Specimens from Morocco, agree with the Etiropean form; 
they have a median dark stripe on the pronotttm, and the long- 
itudinal veins are faintly marked with dark. Also known from 

Creagris murinus (Klug). 

Egypt (Petersen coll.). The median marks on the pronotum 
are in form of two approximate dark lines; the venation is 
wholly yellowish. On Klug's plates it is figured as M. syriacus, 
but in text described as M. murinus. 

Creagris africanus (Rambur). 

From D. O. Afrika, Sansibar, Victoria Nyanza, and D. S. 
W. Afrika (Outyo). The legs are less marked than in C. muri- 
nus, the median stripe on the pronotum is often absent, and the 
wings are plainly more acuminate at tips than in C. murinus. 
Peringuey's species, C. pseudo plumbeus from South Africa 
seems to be the same species. It is probable that this and C. 
murinus are but varieties of C. plumbeus; C. luteipennis, C. 
lanceolatus Rbr., and C. lupinus Oliv., are also closely related 
if not identical, and McLachlan suggests that several of Wal- 
ker's Indian species also belong to C. plumbeus. 

16 Annals Entomological Society of America [Vol. IV, 

Creagris mortifer (Walk.) 

From Daressalam; Reitfontein, D. S. W. Afrika (at light), 
and Abyssinia. Probab y occurs over most of Africa. 

Creagris aegyptiacus (Rambur). 

From Egypt (Petersen coll.) ; Bagamoyo and Korogue, D. O. 
Afrika, also Madagascar. Also widely spread in Africa. 

Peringuey has recently described four other species from 
South Africa. One, C. angustipennis, may belong to the genus, 
but the three other species (C damarinus. C. mashiinus and 
C. bechiianus) are said to have the spurs no longer than the first 
joint of tarsi; since all the species of Creagris known to me have 
spurs as long as three or four joints, I can hardly believe that 
these three species really belong to Creagris. 

Formicaleon n. gen. 

This genus, of the general appearance of Myrmeleon, is at 
once distingiushed therefrom by having but one crossvein 
before the radial sector in the hindwings, and by the antennae 
being close together at base; from the allied genera, Nemoleon 
and Macronemnrus, it is separated by the longer- spurs. 

The genus Formicaleo Leach was made for Myrmeleon for- 
micarius, which was already the type of Myrmeleon; therefore, 
I am compelled to make a new name for the Formicaleo that is 
used by Hagen and Brauer. 

The type is Myrmeleon tetragrammicus Fabr. 

There are many species in the African fauna, the few I 
have seen may be separated as follows: 

1. Two broad black stripes on the yellow pronotum, and the lower sides black, 

antennae long and slender, wings rather broad, most of the cross-veins 

brown 2 

Not two broad black stripes on a yellow pronotum. .' 3 

2. Two cross-veins before radial sector in the hind-wings, stigma rosy, .Idoneus 
But one cross-vein before radial sector in the hind-wings; stigma white. . . 

.3. Hind-wings with a dark streak in apical part; pronotum pale, with incomplete 

dark lines harpalyce 

No streak in the hind-wings 4 

4. Four dark spots just behind the radius in each fore-wing hesione 

No such spots 

-5. Hind part of fore-wrings with many small dark spots; pronotum dark, with 

two darker stripes; antennas short lepidus 

Not so 6 

6. Pronotum with a broad median black stripe, and oblique black line in apical 

part of the fore-wings ilione 

Pronotum and wings not so marked 7 

7. Gradates in fore-w'ings margined with brown; larger species, expanse 80 mm. 

Gradates of fore-wings not margined; smaller species, expanse about 50 mm. 


1911] Notes on African Myrnieleoiiidce 17 

Formicaleon harpalyce n. sp. 

Head yellow, a black mark under each antenna, a large interan- 
tennal mark, mostly above the antennas, two bands across vertex made 
up of spots, a median spot behind and one each side on occiput; antenna; 
yellowish, tip darker, basal joint with dark mark below; pronotum 
broad, yellowish (or reddish), two black marks on front margin, a sub- 
median pair of stripes behind, the posterior side margins broadl}^ behind, 
and a spot each side on the transverse furrow black; rest of thorax 
pale, with black marks, mostly over base of wings, metascutellum with 
a pale median line; abdomen shorter than wings, dull black; legs pale 
reddish }'ellow, spurs curved, the first pair as long as four joints. 
Wings hyaline, ^'enation pale, longitudinal veins not interrupted with 
dark, or very faintly so, some crossveins, especially costals partly dark, 
stigma yellowish (or reddish); hind wings with a long brown streak 
from the anastomosis of cubitus and median outward toward tip of 
wing. Wings slender, acute at tips, hind wings falcate at tip; about 8 
crossveins before radial sector in forewings, 12 or 1.3 branches of radial 
sector in both pairs. 

Expanse 70 to 80 mm. 

From West Africa, Bismarcksburg, Togo. 

Formicaleon alcione n. sp. 

Face pale yellowish, black band below and above antennae, a trans- 
verse frontal band on vertex, and spots more or less connected in a row 
behind; antenna? pale, annulate with brown, ti]3 dark; pronotum broader 
than long, dark, three ill-defined pale stripes, the median more narrow; 
thorax pale, marked with dark, scutelli with a double dark spot; abdomen 
shorter than wings, blackish, pale each side near tip; legs pale, femora 
spotted with black, especially near the tip, tibia with bands above 
near base and tip, tips of tarsi black; the tarsi short, last joint nearly 
as long as other joints together, spurs as long as four joints. Wings 
hyaline, venation black and white, not dotted, but in long streaks, a 
spot at base of stigma, and two spots (smaller) beyond on the radius, 
the outer gradates very irregularly, but plainly dark, and a faint dark 
streak up from end of anal vein; hind wings with stigmal spot, and two 
beyond dark, also one on the anastomosis of cubitus and median. Wings 
very long and slender, acute at tips, hind wings falcate at tip, 7 or S 
crossveins before radial sector in forewings, 14 or 1.") branches of radial 
sector in both pairs. 

Expanse 75 mm. 

From Mohorovi, D. O. Afrika, and one marked "Key. 
Frustorpher. " 

Formicaleon idoneus n. sp. 

Face pale, a large black interantennal mark, reaching below the 
antennas where it is deep and shining, paler above toward vertex, two 
rows of transverse spots on the vertex, the hind row with a projection 
each side behind; antenna; long, pale on base, and annulate with brown, 

18 Annals Entomological Society of America [Vol. IV, 

beyond and at tip darker brown; pronotum broader than long, yellowish, 
with two broad black stripes and the lower margins black; rest of thorax 
pale, striped and spotted with black, scutelH mostly pale, the mark 
over base of the forewings includes two pale sjDots; abdomen shorter 
than wings, brownish, jialer on basal half and there with lateral black 
line; legs pale, anterior femora and tibiae dotted with black, tip of tarsi 
black, apical joint of tarsus nearly as long as others together, spurs about 
as long as four joints. Wings hyaline, stigma rosy, longitudinal veins 
black and white, radial sector and branches and most of the crossveins 
black. Wings rather broad beyond middle, tips acute, hind wings sub- 
falcate; S crossveins before radial sector in forewing, two in hindwing, 
13 branches of radial sector in both wings. 
Expanse <S.5 mm. 

From Langenburg, D. O. Afrika. In general similar to 
F. lethalis, having broad wings and slender antennae, but the 
two crossveins before radial sector in hindwing distinguisli it. 

Formicaleon ilione n. sp. 

Face pale, a vertical black line between bases of antennae; a broad 
deep black band above antenna: ; two, rather curved, bands on the vertex, 
connected in middle by dark, and the posterior one extended behind 
in middle; pronotum with a broad black median stripe, broken up 
behind into lateral projections, a spot each side near front on the trans- 
verse groove, and the ]JOSterior sides black; rest of thorax mostly black, 
with a broad pale stri])c each side, sides of scutelli pale; abdomen black, 
each segment (beyond the basal) with a pale stripe each side, connected in 
front of the dorsum, venter rather pale, the pleura? of thorax heavily 
streaked with black, leg Short, hairy; femora thick, with many white 
hairs and bristles and a few black ones, anterior femora mostly black, 
others paler, especially below, front and middle tibia- with two black 
bands, hardly visible in hind tibiae, tarsal joints tipped with black, very 
short, the basal especially so, the fifth as long as all others together, 
spurs longer than three joints. Wings with the longitudinal veins 
heavily marked with black and white, the costals and many crossveins 
mostly black, black spot at base of stigma, a long curved prominent 
black stripe over the gradates toward tip of wing, and a short, oblique 
mark upward from near the end of the anal vein; hindwings less marked, 
the anal vein wholly, and the median vein on the basal third very pale 
yellowish, other veins marked slightly with black. Wings not very 
long, quite broad at stigma, hind pair much narrower, both pairs acute 
and subfalcate at tip, the outer margin being slightly concave, seven 
cross-veins before origin of the radial sector in the forewings, ten to 
eleven branches of the radial sector in each wing. 

Expanse 60 to (3.5 mm. 

From Harrar, Abyssinia (Kristensen collector), types in 
collections Banks and Petersen. 

1911] Notes on African MyrmeleonidcB 19 

Formicaleon persephone n. sp. 

Face pale yellow, a black band below antennte and one above, two 
transverse rows of spots on the vertex; antennae pale, annulate with 
with brown; pronotum broader than long, dull blackish, a faint median 
pale line and an indistinct outer pale stripe each side; rest of thorax 
dark, marked a little with pale, posterior margin of anterior lobe of 
mesothorax pale; abdomen blackish, a pale spot above before middle of 
each segment; legs pale, hind femora and tibiae with blackish dots, 
front femora darker, tibite with basal and apical dark mark, and tips of 
tarsi black; apical joint of tarsus not as long as the others together, 
spurs as long as four joints, curved. Wings extremely slender, fully 
six times as long as broad, acute at tips; venation mostly black, the 
radius mostly pale, the subcosta interrupted black and white, cubitus 
with longer streaks of black and white, cross-veins on large patches are 
pale, the gradates black, a black spot at base of stigma and over union 
of subcosta and radius; 7 crossveins before radial sector in forewings, 
about 11 branches of radial sector in each wing. 

Expanse 50 mm. 

From Thies, Senegal. Its small size and very narrow wings 
separate it from all others. 

Formicaleon hesione n. sp. 

Face pale, a black band below and one above antennae, pale between, 
vertex brown, three double black spots behind in a transverse row, and 
in front of these are two others, submedian; antennas pale, annulate with 
brown; pronotum broader than long, dark, a median pale line, and a 
large anterior spot each side, not clearly outlined, but reaching backward; 
rest of thorax dark, marked with pale, especially on the scutelli; legs 
short, pale, tibije marked with brown above near base and at tip, tips 
of tarsi dark, dark dots on hind femora, black hairs below on femora 
and tibiae I, tarsi short, apical joint about as long as others together, 
spurs as long as four joints, curved; abdomen shorter than wings, 
blackish, a pale spot above before the middle of each segment, apical 
segment with a pale spot each side near tip. Wings hyaline, venation 
black and white, some large patches of white veins, costals black at 
each end, four spots behind radius, one before radial sector, one on origin 
of radial sector, one on first branch of radial sector, and one near stigma, 
the stigma with basal black mark, and a spot on radius beyond, a short 
oblique mark over end of anal vein, and the outer gradates plainly 
black, also a spot over the first crossvein between cubitus and cubital 
fork ; in hind wings a spot at base of stigma, one just behind radius near 
by, and one on radius beyond stigma. Wings very slender, acute at 
tips, 7 cross veins before radial sector in forewings, about ID branches 
of radial sector in both pairs. 

Expanse 5S mm. 

From Nssanakang, Kamerun. In general appearance it is 
very much like Nemoleon ^-maculatus from East Africa, but the 
gradates are more plainly marked, and the tarsi are very 
different in structure. 

20 Annals Entomological Society of America [Vol. IV, 

Formicaleon lethalis (Walker.) 

Specimens from Bagamoyo, Madagascar, Natal, and Mhon- 
ta, D. O. Afrika. Recorded from various places in East Africa. 

The F. leiicospilos Hagen is the same species. 

Formicaleon lepidus (Kolbe). 

From Langenburg, D. O. Afrika, and Erythrsea, (Petersen 


Nemoleon Navas. 

This genus has but one crossvein before the radial sector 
in the hindwings; the antennae are not their diameter apart "at 
base; the first joint of tarsus is about as long as the last joint, 
and the spurs about as long as this first tarsal joint. The wings 
are slender, with a single costal series. 

Type — Myrmeleon arenariiis. 

1. Wings with some dark spots ,3 

Wings without dark .spots '- 

2. Spurs and claws black. kituanus 
Spurs and claws reddish filiformis 

3. Outer gradates in fore-wings form a prominent oblique dark stripe pardalice 
Outer gradates do not form a prominent stripe 4 

4. Four black spots just behind the radius in the fore-wings, and one at tlic 

stigma quadrimaculatus 

A black dot at end of anal vein in fore-wings, and one near end of median 
where it joins the cubitus alcidice 

Nemoleon alcidice n. sp. 

Face pale, a dark brown band over base of antennae, and two nar- 
row brown bands on the vertex, made up of conjoined spots; antenna 
pale, the second joint dark above, the tip also dark; pronotum pale, a 
brown spot on each side margin; and two black dots on each side, one 
on the transverse groove, the other behind it on the hind margin; 
rest of thorax dark on sides, but only faintly marked in the middle, not 
plainly on the scutelli; abdomen dull black, last segment slightly paler 
at tip^ all white haired. Legs slender, pale, distinctly dotted with black 
on the femora and tibias, and at tips of tarsal joints, claws long, reddish, 
as also the spurs, which are as long as the first tarsal joint, and this as 
long as the next three together, apical joint nearly as long as the basal, 
both black and white bristles on the legs. Wings hyaHne; venation 
pale, interrtiptcd with black, fore-wings with an oblique black mark 
near end of anal vein, a roundish, black spot near end of median at its 
connection with the cubittts, one black spot on the union of subcostal 
and radius, and two beyond on the radius, the outer one the larger; 
the anal vein pale, but its branches marked with black on the margin. 
Wings long and slender, acute at tips ; seven crossveins before the radial 
.sector in forewing, nine branches of the radial sector in each wing. 
Hindwings mostly unmarked. 

Expanse .o.S mm. 

From Erythrasa (Kristensen coll.). Type in coll. Petersen; 
(cotype in Banks coll.); a poor specimen from Duma, Kamerun, 
Jtme (Berlin Museum). 

1911] Notes on African Myrmeleonidce 21 

Nemoleon pardalice n. sp. 

Face ]3ale, a dark band under and between antennse, two narrow 
bands on the vertex, made up of black spots, and behind a pair of sub- 
median marks; antennae brown, except the pale base; pronotum dark, 
with a pale spot each side in front, and a pale streak behind; thorax 
dark, a pale spot each side on the anterior lobes, and on scutelli, posterior 
margin of the anterior lobe and of the mesothorax pale, two pale marks 
on the metanotum; femora mostly dark, but white-haired, tibirs with 
basal and preapical dark bands, and tips of tarsal joints dark; spurs 
not as long as the basal joint, which is about as long as the next two 
together, apical a little longer than the basal. Abdomen black, white- 
haired, tip pale. Wings hardly hyaline; venation mostly dark, the 
subcosta, radius, and cubitus, marked with white, some crossveins white, 
and some white on branches of the radial sector, median vein wholly 
black; many veins bordered with black, especially the crossveins in 
hind part of wings, also the outer gradates, which form a long oblique 
line, and the veinlets obliquely above the end of the anal vein, the 
origin of the radial sector, an oblique vein before it, one oblique vein con- 
necting it to the radius, and where it bends up toward the radius near 
the stigma, also black; the stigma dark; some small cross-veins beyond 
the stigma in the costal area are bordered with black ; in the hindwings 
the subcosta black and white, the other veins mostly black, the forks 
of veinlets near the outer hind margin are dark, and also above the 
radius near tip of wing with dark spots on the forks. Wings rather 
short, and broad; forewings hardly acute at tips, hindwings plainly so; 
six crossveins before origin of radial sector in the forewing, one of them 
oblique, about eight branches of radial sector in both pairs, all costals 
simple. Abdomen much shorter than wings; vertex high and con^^ex 
in a ridge, pronotum about as long as broad. 

Expanse 40 mm. 

From Erythraea (coll. Kristensen), type in collection Peter- 
sen. • 

Nemoleon 4-maculatus n. sp. 

Head pale, a black mark each side under antenna along the side of 
the eyes, a black band above antenna;, emarginate above in middle, 
two rows of black spots on vertex, six in each row, more or less connected 
in pairs; antennae pale, annulate with brown; pronotum slender, nar- 
rowed in front, dull black, a median pale stripe from before middle to 
tip, and a stripe each side pale; rest of thorax dull black, with a few 
yellowish spots; abdomen about as long as wings, black, with white 
hair, tips of segments with a yellow spot above; legs pale, marked with 
black, long black bristles and white hairs, tips of tibiae and tarsi black, 
femora with black near middle, at least above; basal joint of tarsus as 
long as last, spurs a little longer than first joint. Wings hyaline, marked 
with black, venation black and white, forewings with four black spots 
behind radius, one before origin of radial sector, one on origin of radial 
sector, one a little beyond first branch of radial sector, and one near the 

22 Annals Entomological Society of America [Vol. IV, 

stigma; base of stigma black, a narrow streak up from end of anal \-cin, 
spot at anastomosis of cubitus and median, and two dots on radius 
beyond stigma; many crossveins are black, but there are large patches 
where all are white, two of these near middle of wing and one oblique 
below the stigma; hind wings with spot at base of stigma, and a spot 
behind radius black, some of the same large patches of white crossveins 
as in the forewings. Wings very slender, acute at tips, 7 crossveins 
before radial sector in forewings, about 9 or 10 branches of radial sector 
in both pairs. 

Expanse oS mm. 

One from Langenburg, Nyassa-See, and one from Zanzibar. 

Nemoleon filiformis (Gerst.). 

From Bismarckburg, Togo, and Nssanakand, Kamerun. 

Nemoleon kituanus (Kolbc). 

From Peleki, Kwidjwi, D. O. Afrika. ■ 

Macronemurus Costa. 

This genus is similar to Forniicaleon, but the spurs are 
shorter. The antennae are close together at base ; the pronotum 
is very narrow when compared with the head ; and the antennae 
are rather long and slender. There are two types of venation; 
one in which the forks of cubitus are much divergent, and the 
wings are broader, and the radial sector arises barely beyond the 
forking of cubitus. This is the typical section. The other 
group has narrower wings, the radial sector arises much beyond 
the forking of cubittis, and the cubital forks are nearly parallel 
approaching the condition of Creagris; the species M. iolanthe, 
and M. melanthe described below belong to this section. 

Type of genus — M. appendiculatits Costa. 

The species here recorded may be separated by the following 
table : 

1. Hind-wings with an apical brown streak; cross-veins all dark; pronotum with 

dark stripes striola 

Hind-wings with the outer apical margin dark; a dot at end of anal and on 

anastomosis of cubitus and median in the fore-wings iolanthe 

Hind wings without streak or dark margin 2 

2. Fore-wings with many large black spots (about 24) and four or five near 

apex of the hind-wings pulchellus 

Wings without large marks 3 

3. Radius and subcosta unmarked; cross-veins nearly all pale; no spots on wings; 

pronotum mostly dark melanthe 

Radius and subcosta black and pale alternately 4 

4. Pronotum pale, with three entire, black stripes; gradate veinlets marked; 

many cross-veins are margined with brown at their junction with the 

longitudinal veins loranthe 

Pronotum not so marked 5 

1911] Notes_ on African Myrmel eon idee 23 

5. All the cross-veins are margined with brown; pronotum pale, with lateral 

dark stripe, and three dark spots in the middle ianthe 

All cross-veins not margined with brown 6 

6. A median forked stripe on pronotum; cross- veins nearly all dark; stigma 

white tinctus 

No median forked stripe on the pronotum 7 

7. Pronotum yellow, with two spots at front margin and a median one behind, 

and a narrow stripe each side dark; nearly all cross-veins dark . . .euanthe 
Pronotum mostly dark S 

8. Pronotum dark with a narrow, pale, median line chloranthe 

Not so marked 9 

9. A black dot at end of anal vein, and one at base of stigma; pronotum all 

dark (a broken specimen seen) sp. 

No dots on wings; pronotum marked with pale, (a broken specimen seen) . .sp. 

Macronemurus striola Kolbe. 

From Bagamoyo, and Lindi, D. O. Afrika; the former 
specimen is small, and slightly marked, but does not seem to 
differ in structure. 

Macronemurus tinctus Kolbe. 

From Peleki and Netron See, D. O. Afrika, and Kiiui, 
East Africa. 

Macronemurus chloranthe n. sp. 

Face yellow, a black band below and a broader one above antennae, 
two rows of dots on the vertex, the hind one cur\'ed; antennte slender, 
broadly annulate with brown; pronotum fidly as long as broad, sides 
subparallcl, dark, a narrow, median pale line, and a pale sinuate stripe 
each side, a black dot each side on the transverse furrow, thorax dark, 
black on middle of metanotum, and some faint pale marks in front; 
abdomen blackish, a pale yellowish streak each side above on middle 
of each segiuent, shorter than wings; legs jjale, femora darker near tip, 
tibiae with subbasal and apical marks brown, joints 2, 3, 4, and apical 
part of 5 black, legs white-haired, a few black bristles, apical joint of 
tarsus much longer than broad, sjjurs fully as long as two joints. Wings 
hyaline, venation black and white, usually in long streaks, especially 
prominent on subcosta, radius, and cubittxs, a dark spot at base of stigma, 
and the crossvein below it black, outer gradates dark and spot at end 
of anal vein; in hind wings with spot at stigma; wings slender, acute at 
tips, 7 crossveins before radial sector in forewings, 8 branches to radial 
sector in both pairs. 

Expanse 4() mm. 

From Okahanda, S. W. Afrika. 

Macronemurus iolanthe n. sp. 

Face yellow, interantennal mark and the vertex all black; antennae 
long, brown, tip darker; pronotum a little longer than broad, narrowed 
in front, dull blackish, with faint median pale liiie, and a broader pale 
mark on each side, indistinct in some lights; thorax and abdomen black, 
latter with white hairs, apical segment pale; abdomen of female shorter 
than wings; legs with rnostly black femora, anterior pairs paler in front, 

24 Annals Entomological Society of America [Vol. IV, 

tibias I and II blackish above, tarsi long, but apical joint much longer 
than basal, spurs about as long as two joints. Wings hyaline, outer 
margin near tip fuinose, more prominent in hind wings, a spot on fork 
of cubitus, one below stigma, two on radius beyond stigma, one at 
anastomosis of cubitus and median, and an oblique one at end of anal 
vein, black, some small dots along anal ^^ein beyond its middle; hind 
wings unmarked, \-enation mostly pale, the subcosta, radius, and 
cubitus faintly dotted with dark. Wings long and slender, acute at 
tips, 7 crossveins before radial sector, 10 or 11 branches of radial sector 
in both pairs; the anal (in forewings) runs parallel to cubital fork for 
some distance, six or seven cross-veins between them. 
Expanse 04 mm. 

From Bismarckburg, Togo. 

Macronemurus melanthe n. sp. 

Face yellowish, interantennal mark and vertex entirely black; 
antennae brown, darker at tips; pronotum a little broader than long, dull 
\-cllowish brown, with indistinct dark marks, a pair in front, a pair 
behind, and the outer posterior edges black; thorax mostly black; 
abdomen blackish; legs pale, femora and tibife dotted with red brown, 
tarsi long and slender, last joint as long as the first three joints, spurs 
only a little longer than basal joint. Wings hyaHne, venation pale, 
almost unmarked, stigma not distinct. Wings long and slender, acute 
at tips, 6 crossA'cins before radial sector in forewings, 11 branches of 
radial sector in both pairs ; in forewings the anal vein inns jjarallcl to the 
cubital fork farther than in other species (except iolanthe), there being 
five to seven cross\'eins between them. Male abdomen is longer than 
the wings, the appendages rather long, but not one-half as long as the 
penultimate joint. 

Expanse 58 mm. 

From Bismarckburg, Togo. 

Macronemurus euanthe n. sp. 

Head yellow, a black band above and below antennas, a narrow trans- 
verse line across front of vertex, and a curved row of six small dark 
spots behind, behind this a median spot; antennae rather long, pale, 
faintly annulate with brown, pronotum not much broader than long, 
pale yellowish, two submedian si)ots on front margin, a median spot 
iDehind, and a stripe each side, brown, the latter not reaching front 
margin; thorax yellow, spotted with dark, anterior lobe with two spots 
and the anterior margin dark, scutelli with median dark stripe, and dark 
streak over base of wings ; abdomen shorter than wings, dark, pale on base, 
white-haired; legs pale, some dots on the hind femora, a sub-basal and 
apical mark abo\'e on tibiffi I and II, and tip of tarsus dark, long erect 
bristles on femora and tibiae, last joint of tarsus nearly as long as the 
others together, spurs as long as three joints (leg I) or two joints (leg 
III). Wings hyaline, nearly all crossveins dark, and margined with 
pale brown, longitudinal veins pale, the radius and cubitus with dark 
dots; in hind wings the cross\'eins not margined, but some dark dots 

1911] Notes on African Myrmeleonidce 25 

near apical margin. Wings rather slender, forewings acute at tips, hind 
wings subfalcate; 7 cross veins before radial sector in forewings, 9 
branches of radial sector in both pairs. Male abdomen longer than wings 
very slender, appendages filifonn, about as long as the penultimate 

Expanse 45 mm. 

From Reitfontein, D. S. W. Afrika, (at light). 

Macronemurus ianthe n. sp. 

V^ery similar to M. euanthe, the same spotting of pronotum and 
thorax ; the head has the interantennal mark reaching farther up on the 
vertex so there is no hne across the vertex and in place of the row of 
spots is an anchor shaped mark, and a dot each side adjoining the eye; 
abdomen yellowish brown, white-haired; legs as in M. euanthe, the spurs 
hardly longer than two joints; wings as in M. euanthe, but all cross-veins 
are more heavily bordered with brown, the subcosta, radius, and cubitus 
are prominently white and black; in hind wings many crossveins, esjjec- 
ially in the posterior part of wings, are margined with brown, the sub- 
costa, radius, and cubitus white and brown; stigma yellowish. Wings 
slender acute at tips, 7 crossveins before radial sector in forewings, 10 
branches of radial sector in both pairs. 

Expanse 56 mm. 

From Langenburg, D. O. Afrika. 

Macronemurus loranthe n. sp. 

Head yellow, a large black interantennal mark extending below 
antenna and much above, and on middle above it is connected to a me- 
dian dark stripe which extends to the back of head, a brown spot each 
side on vertex; antennae brown; i^ronotimi much broader than long, 
yellow, a broad median and a lateral stripe each side dark brown, each 
■continued back on the mesothorax, the lateral ones furcate behind; a 
median stripe on metanotum, and a furcate mark over base of wings; 
pleura with a broad interruiJted brown streak. Abdomen slender, in 
male longer than wings, and with appendages nearly as long as last 
segment, pale brown, a faint pale mark each side near base of segments, 
sometimes in form of a streak. Legs pale yellow, the tibias with basal 
and apical dark spot above, tip of tarsus dark, and dark dots on femur, 
last tarsal joint nearly as long as the others together, the spurs nearly 
as long as three joints. Wings hyaline, venation pale, marked with 
brown at connection of all veins, thus making many small dots all over 
the wing, more prominent along ctibitus and on outer gradates, stigma 
pale, no basal spot; hind wing with these small spots along the hind and 
apical margins; the subcosta and radius interrupted with brown. Wings 
rather broad at stigma, acute at tips, hind wings sub-falcate at tip; 
5 to 7 crossveins before radial sector in forewing, 12 branches to radial 
sector in both pairs, in forewings a few of the costals before stigma 
are forked. 

Expanse 50 to 57 mm. 

From Bismarckburg, Togo, and the Togo Hinterland. 

26 Annals Entomological Society of America [Vol. IV,. 

Macronemurus pulchellus n. sp. 

Face pale, a black dot between antenna, a dark band above antcnnje, 
and a spot each side on posterior part of vertex; antennae pale brownish; 
pronotum yellowish, unmarked, except a black dot each side on hind 
margin and a black mark in posterior comers ; the outer part of pronotum 
before the groove is rather darker than the middle; rest of thorax pale 
yellow, black on sides, and across the anterior lobe, two spots on the 
mctathorax black, scutelli wholly pale; abdomen black, segments 
broadly pale behind, tip and most of the second segment pale; legs long 
and slender, hind femora deep-black, tibia lined near the tip below, 
anterior femora with an outer black line, spurs as long as two tarsal 
joints, the basal joint very short, the apical nearly as long as all others 
together. Wings hyaline; forewings heavily maculate with many quite 
large black marks, and some near the apex of the hindwings ; costal area 
of forewings unmarked, about 24 spots over rest of surface, mostly over 
a crossvein, and roundish or elongate in shape; the elongate ones mostly 
near and at right angles to the hind margin; hind-wings with four or 
five such spots near the apex and toward hind margin; venation mostly 
pale; stigma not very dark. Wings rather slender, acute at tip; fore- 
wing with about five crossvcins before radial sector, nine branches to 
radial sector in each wing; abdomen shorter than wings. 

Expanse 65 mm. 

From Erythrjea (Kristensen coll.); type in coll. Petersen. 

Cymothales Gerst. 

The radial sector of hind wings arises near base, with but 
one crossvein before it ; the wings are rather broad in the stigmal 
region, and more or less falcate at tip, the hind pair usually 
strongly falcate; the cubitus in hind wings is scarcely forked, or 
the fork so weak as not differing from the other veinlets from 
cubitus to the margin; the anal also weak. The antenna 
are very slender, and rather wide apart at base; the legs are 
extremely slender, the basal tarsal joint longer than the apical; 
spurs as long as two joints; the tibise longer than the femora; 
the prothorax is long and slender. 

Type — C. mirabilis Gerst. 

The species can be separated by this table: 

1. Middle band of fore-wings plainly divided into two spots 2' 

Middle band of fore- wings undivided, at least one connection 4 

2. This band only narrowly divided; the apical mark of fore-wings is very broad, 

and hardly at all broken up by pale spots liberiensis 

This band is broadly divided: the apical mark broken up by pale spots. ... 3 

3. Femur I black, and black-haired; antennae mostly pale; a spot on fore-wings 

before the stigma, and one near middle of hind margin of each wing. . . . 

Femur I not black-haired; and only with a few pale dots eccentros 

1911] Notes on African MyrmeleonidcB 27 

4. Thorax black, with narrow pale lines 5 

Thorax pale, with a median black stripe over meso and metathorax; the spot 

on hind margin of hind wings is connected along the margin to the apical 
spot delicatus 

5. Antennae pale, except basal joint and the tip dulcis 

Antennce dark for one- third way out (i 

6. Continental specimens mlrabilis 

Madagascar specimens bouvieri 

Cymothales liberiensis Wcclc. 

From Longji and Victoria, Kamerun; I have one from 
Sierra Leone. 

Cymothales mirabilis Gcrst. 

From Bismarckburg, Togo, and Abyssinia (Petersen coll.) 
Known from several other localities in Africa. 

Cymothales speciosus Gerst. 

One from Amani, D. O. Afrika. 

Cymothales sp. 

A broken specimen from Kongo, appears to be a new species 
related to C. delicatus. 

Cymothales delicatus n. sp. 

Face pale, a curved brown mark on each side, a band below antenna?, 
above brown and on the front part of the elevated portion, vertex with 
six brown spots in a transverse row in front, and three larger sijots in a 
row behind, each of the latter are nearly double; antenna with dark 
mark on basal joint, and a dot each side on the second joint, beyond 
black for about five or six joints, and then white for three joints, then 
brown to the tip; pronotum pale with a median stripe brown, and a 
brown curved line each side in front; rest of thorax dark brown, with a 
white stripe each side; abdomen pale brown; legs pale, femora mostly 
brown, especially above, tibia* with brown dots. Wings hyaline, vena- 
tion mostly pale, but the veins in dark spots are dark; forewings with 
a basal spot, with an outer extension behind; a median spot from 
radius to the median vein nearly square, and narrowly connected to a 
brown V-mark behind it; a long brown stigmal mark, extending obliquely 
inward, not so well defined, to near the median vein; slightly behind it 
begins an oblique, brown streak which extends to the hind margin and 
in front points toward the tip of the wing; around the tip are several 
dark spots, but not very distinct, the whole apical area is slightly fumosc; 
the costal area has several of the crossveins marked with brown. Hind 
wings with a basal brown spot over origin of the radial sector, one on a 
crossvein from radius to radial sector, about seven crossveins beyond the 
basal one, another toward the stigma, and one under stigma, the apical 
part above radius mostly dark, but a narrow apical pale streak, hind 
margin toward tip has two well separated brown spots, the outer one is 
connected by brown along the hind margin to the brown at tip, the 

28 Annals Entomological Society of America [Vol. IV, 

short veinlets along the hind margin arc mostly dark. Wings broad; 
the forewings hardly falcate at tip; the hind pair but little longer than 
front wings, falcate at tip, but not as strongly so as in C. mirabilis; 
about four crossveins before the radial sector in the forewings, about 
eight branches to radial sector in each wing, the sixth branch is soon 
forked, otherwise venation much as in other species. 
Expanse 68 mm. 

From Kongo, West Africa, in coll. Banks. 
Echthromyrmex McLach. 
(Palparidius Peringuey.) 

Antennae hardly diameter of basal joint apart, not very long; 
legs rather short, basal joint short, apical as long as others 
together, the sptirs nearly as long as two joints, tibiae shorter 
than femora; pronotum very short; male appendages extremely 
long. Wings rather broad; costals simple, one crossvein before 
radial sector in the hind wing, in forewings the first branch of 
radial sector arises barely before the cubital fork. In hind wing 
the cubital bends upward and then downward to the anasto- 
mosis with the median, the anal parallels it so that the cubital 
fork is indistinct. 

Type — E. platypteriis McLach. 

I have not seen the type species, and the above description 
is taken from the species described below which I believe is 
congeneric; at least it agrees with McLachlan's description, and 
is remarkable in having, as he said, the "postcosta simple" 
(= cubitus); in this feature it is related to the South American 
genus Dimares, and the markings of the wings are also similar 
to that genus. The genus, Palparidius, recently described by 
Peringuey is the same, without doubt ; it has no very close 
relation to Palpares. 

Echthromyrmex fascipennis n. sp. 

Face yellowish; a transverse brown mark on face, a large brown 
interantennal mark slightly pointed below on the middle, and extending 
up in front of the vertex, which is bilobed; four pale brown marks on 
vertex in a curved row; antennae dark brown; pronotum very short, 
nearly twice as broad behind as long, much narrowed in front, pale, 
with a median black stripe, and an oblique lateral one each side, many 
minute black points at base of bristles; rest of thorax black, a broad 
pale stripe over middle behind the anterior lobe, and including a median 
dark stripe; thorax at base of wings pale; abdomen dark, a pale median 
and a lower lateral stripe, all very narrow toward tip; appendages yel- 
lowish, blackish near base below; legs yellowish, rather short and heavy, 
not very hairy, basal tarsal joint barely longer than the second, apical 
as long as others together, spurs straight, nearly as long as two joints. 

inil] Noles oil African Myrniclconidcc 2!) 

Wings h3'alinc, \'\-itli large black marks; forcwings with dots along the 
upper edge of the subcosta along both sides of the cubitus, to the fork, 
and along the lower branch of the fork; many dots near the posterior 
and outer margin; two oblique narrow black "bands, both begin at the 
radius, one before the middle and one beyond the middle, this latter is 
broken near tip, and reaches in a line to the hind margin; an oblique 
apical streak black, and including several white crossveins, a dark mark 
before and one beyond the white stigma. Hind wings with some dots 
along the subcosta, a rather broad oblique dark band before the middle, 
beginning at the radivis and with a curved tip, not reaching across wing; 
a large dark spot before the stigma, and behind it on the hind margin are 
two narrow upward streaks, the inner one forked, a short apical dark 
streak, and some dark dots along the outer margin; venation of both 
pairs mostly pale, except dark spots on the radial sector and cubitus. 
Wings rather broad, not very long, not acute at tips; three or four cross- 
veins before radial sector in forewing, about nine branches to radial 
sector in each wing, the anal region of both pairs is very broad, in the 
hind wing the post-anal extends ovit quite a- long distance, and has five 
branches to the margin. Abdomen shorter than the wings (in the male), 
the male appendages are extremely long, fully one-half as long as tht; 
abdomen, eur\'ed downward, and parallel. 
Expanse 7() mm. 

From Okahanda, Deutsch Sud-West Afrika, (Berlin Mu- 
seum) . 

Peringuey has recently described* two species which are 
closely related to E. fascipennis. He made for them a new gemis, 
Palparidius, but his figure shows it is the same as Echthromyr- 
mex. His P. capicola has the stigmal mark of the hind wings 
entire, not broken up into three spots as in E. fascipennis; 
his P. concinnus has this stigmal mark broken up, but the median 
band is very narrow, and not hooked at end; there are several 
other differences in maculation. The male appendages of E. 
fascipennis appear to be much longer than in either of Perin- 
guey's species. 

Nemoleon pardalicc. 
Formicaleon ilione. 
Myrmeleon medialis. 
Nesoleon boschimanus. 
Nesoleon punctatissimus. 
Creagris pretiosa. 
Macronemurus tnelanthe. 
Nemoleon quadrimaculatus. 
Echthromyrmex fascipennis. 
Echthromyrmex fascipennis. 
Macronemurus loranthe. 

* Annals South African Museum, V, p. 43, 1910. 






Myrmeleon alcestris. 





Macronemurus ianthe. 





Myrmeleon furcatus. 





Gymnoleon exilis. 





Myrmeleon doralice. 





Nemoleon alcidice. 





Myrmeleon obscurus. 





Formicaleon harpalyce. 





Macronemurus euanthe. 





Macronemurus chloranthe. 





Formicaleon ideonus. 





Macronemurus loranthe. 

Annals E. S. A. 

Vol. IV, Plate I. 

Nathan Bittd-i^. 


Vol. I\', Plate II. 

Nathan Banks. 



By Ralph V, Chwiberux. 
Provo, Utah. 

In this paper is summarized our present knowledge of the 
genera and species of the Lithobiomorpha occurring in the 
extensive region lying south from Kentucky and the Virginias 
and east of the Mississippi river. The great majority of the 
records given are based upon collections made by the author 
himself in the summer of 1910, during which season all 
of the states in the territory indicated, excepting Florida, 
were visited. The southern portion of Georgia and the coastal 
region of this state and of the Carolinas were not covered. 
Some species additional to those here listed are likely to be 
found in these sections. The season was favorable for mem- 
bers of this group; and it is felt that the great majority of the 
more widespread forms were secured. In this connection it 
may be noted that in the case of most of the species specimens 
were taken in a considerable number of localities and that all 
the species previously recorded were again found excepting 
two from southern Georgia and Florida, where, as before 
mentioned, collections were not made. 

Of especial interest are the genera Buethobius and Wato- 
bius, here erected for the first time, and Zygethobius, previously 
established by the author for a species occurring in the high 
mountain ranges of the western Unites States. The finding 
of a second species of Zygethobius in the mountainous section 
of this southeastern region fulfills what had been anticipated 
as likely. The three genera named are annectant and must 
alter to a considerable degree some prevalent conceptions 
as to affinities within the suborder. The genera recognized 
in the present paper may be separated as follows: 

a. Legs bearing bristles only, no articular spines present excepting sometimes one 
at distal end of tibia of all but last pairs of legs but this usually re- 
placed by an acutely pointed process. 
b. First leg-bearing segment with a pair of spiracles. 

c. A single pair of ocelli; tarsi of first thirteen pairs of legs undivided, 
those of the last two pairs biarticulate. . . .Lamyctes Meinert. 
bb. First leg-bearing segment without spiracles. 

c. Tarsi of first thirteen pairs of legs undivided, those of the last 

two pairs biarticulate; ocelli none 

Buethobius gen. nov. 


inil] Lithohionwrpha of Southeastern States 33 

cc. Tarsi of all legs biarticulate; ocelli present. 

d. A single pair of ocelli present; an acutely pointed process 

at distal end of tibia; reproduction seemingly partheno- 

genetic, no males occurring . . Zygethobius Chamberlin. 

dd. A number of pairs of ocelli present, forming a patch on 

each side of the head; a spine at distal end of tibia; 

males occurring Watobius gen.nov. 

aa. Legs provided with articular spines as well as with bristles; no acute process 
at distal end of tibia on cephalic side. 

b. Coxal pores in a single series Lithobius Leach. 

bb. Co.\al pores scattered or in several series Bothropolys Wood. 

Genus Lamyctes Meinert. 

1. Lamyctes tivius sp. nov. 

Slender, widest at tenth dorsal plate, very gradually attentiated 
cephalad, more abruptly caudad. 

Dorsum yellow to light brown, the head, prehensorial feet and ulti- 
mate segments darker; antennae and legs yellow. 

Antennae of moderate length, composed of twenty-eight to thirty- 
one articles; first two articles long, the third and fourth very shorr, the 
fifth and sixth longer, the seventh and eighth again very short, the ninth 
longer, the tenth and ele\'enth in turn shorter, the twelfth and subse- 
quent articles comparable to the ninth, or the thirteenth and fourteenth 
in some reduced; this alternation of pairs of shorter articles with longer 
ones in proximal portion of antennae apparently constant in this species. 

A single pair of large ocelli. 

Prosternal teeth 2-|-2, small, or 3-|-3, the outer one on each side 

Angles of none of the dorsal plates produced. 

Coxal pores small, round, 2, 3, 3, 3. 

Anal legs long and slender, the joints of tarsus especially so; pre- 
femur long, clearh' more slender proximally than that of the penult 
pair, clavately enlarged distad; tibia of nearly unifonn diameter 
throughout length, the first tarsal joint of similar shape and length 
but more slender. (See PI. 3, fig. 2 cf. also figs. 1 and 3). 

Claw of gonopods entire. Basal spines 2-|-2, ratSier stout, the inner 
considerably smaller. 

Length 6-7.5 mm. 

Localities. — Byram and Holly Springs, Miss.; New Orleans, 
La.; Jackson, Ala.; Atlanta, Ga. ; Hot Springs, N. C. 

2. Lamyctes tivius var. pius, var. nov. 

Agreeing in general with the species as above described 
but conspicuously longer and more robust, the length of spec- 
imens examined lying between 9 and 9.5 mm. Color uniformly 

Locality. — Hot Springs, N. C. 

34 Amials Entomological Society of America [Vol. IV, 

Genus Buethobius gen. nov. 

First leg-bearing segment without spiracles. 

Ocelli none. 

Tarsi of the first thirteen pairs of legs undivided, those of the 
fourteenth and fifteenth pairs biarticulate. 

Legs without true spines. Tibiae of the first thirteen pairs of legs 
with an acutely pointed process at distal end on cephalic side like that 
of Lamyctes and Zygethobius. (See PL 4, fig. 1). 

Apparently only females found and the reproduction 

Type. — Buethobius oabitus sp. nov. 

3. Buethobius oabitus sp. nov. 

General color yellow or light yellowish brown; the head and pre- 
hensorial feet and in some the ultimate segments clear orange ; antennae 
and legs clear yellow. 

Rather slender, for most of length parallel sided. Narrowed over 
a few segments behind head and more abruptly at caudal end. 

Antennae long, reaching the ninth body segment; composed of 
thirty-six articles of which those beyond the second are rather short, 

Prostemal margin wide; teeth 3+3, very small. 

Ocelli none. 

Angles of none of the dorsal plates produced. 

Coxal pores round, moderate in size; 3, 3, 3, 3 — 4, 4, 4, 4. 

Legs of the first thirteen pairs each ending in three claws; those of 
the fourteenth and fifteenth pairs with the claws single. Anal legs long 
and slender. 

Claws of gonopods undivided; basal spines 2-|-2. 

Length 10-12 mm.; width at the tenth plate 1.3-1.4 mm. 

Locality. — Byram and Canton, Miss. 

Genus Zygethobius Chamberlin. 
Sub-genus Zantethobius subgen. nov. 
Angles of the sixth. se\'enth, ninth, eleventh and thirteenth dorsal 
plates produced. 

Type. — Zygethobius pontis, sp. nov. 

The previously described species, Z. dolichopus Chamb., 
the type of the genus, may be placed in a subgenus Zygethobius 
sens. str. 

4. Zygethobius pontis sp. nov. 

Moderately robust; strongly narrowed caudad and cephalad of the 
tenth dorsal plate, the first leg-bearing segment especially narrow. 

Dorsum in color somewhat chestnut, wnth a narrow median longitu- 
dinal stripe blackish, the first segment darker than the others; head 
.deep to blackish brown; prehensorial feet and antennae reddish, the 

1911] Lithobionwrpha of Southeastern States 35 

latter becoming paler, }-elIowish, distad; venter yellowish to light 
brown, the caudal plates reddish; legs brown, paler proximally than dis- 
tally; last pairs of legs darker, blackish proximally, pale distad. 

Antennae very long; composed of forty-three articles which are 

The ocelli of the single pair very large. 

Prostemal teeth 3+3. 

Angles of the sixth, seventh, ninth, eleventh and thirteenth dorsal 
plates produced. 

Coxal pores 3, 4, 4, 4. 

The process at distal end of anterior pairs of legs apically acutely 
spinesccnt. (See PI. 4, fig. 2). 

Anal legs long and slender. 

Claw of gonopods entire; basal spines 2+2. 

Length ad 10.5 mm.; width of tenth plate 1.(3 mm. 

Localities. — Johnson City, Tenn.; Natural Bridge, Va. 

Genus Watobius gen. nov. 

First leg-bearing segment without spiracles. 

A number of pairs of ocelli present, these forming a patch on each 
side of the head as in Lithobius, a caudal one in the place of the so-called 
single ocellus in the latter genus and the others toward the base of the 

Tarsi of all legs biarticulate. 

Legs without true spines excepting one at distal end of tibia of 
anterior legs in place occupied by the process in the preceding genera. 

Fifth joint in penult legs of male greatly enlarged, the anal (in type) 
not modified. 

Both sexes occurring. 

Type. — Watobius anderisus sp. nov. 

5. Watobius anderisus sp. nov. 

vSlender, attenuated from the tenth dorsal plate cephalad, more 
abruptly caudad. 

Brown, the uhiiuate segments often darker; head and prehensorial 
feet conspicuously darker, deep brown or brownish black; antennae 
brown, pale distad; legs light brown, the posterior pairs having a pur- 
plish tinge, the anal and penult pairs abruptly pale distad of the femur, 
the tibiae in the penult legs especially pale. 

Angles of the ninth, eleventh and thirteenth dorsal plates i^roduced. 

Antennae short; in most composed of twenty-two articles, in some 
of but twenty; articles decreasing in length gradually and uniform!}' 
from the first to the penult. 

Ocelli on each side composed of one large posterior one, in place of 
the single one in Zygethobius, etc., and of eight smaller ones in a patch 
arranged in three series; thus, 1+3, 3, 2. 

Prostemal teeth 2+2. 

Angles of the ninth, eleventh and thirteenth dorsal plates produced. 

36 Annals Entomological Society of America [Vol. IV, 

Coxal pores small, 2, 2, 2, 2. 

All legs with three claws excepting those of the ultimate pair \\-hich 
seem to have the exterior accessory claw but to lack the inner one. 

Anal legs of male moderately and unifonnly crassate; the penult 
legs with the fifth joint strongly enlarged and somewhat flattened dorso- 
ventrally, complanate above or weakly depressed, complanate and weak- 
ly furrowed beneath. (See PI. 3, figs. 4 and .5). 

Claw of female gonopods tripartite; basal spines 2 + 2. 

Length 7. .5-9. 5 mm. 

Localities. — Thomasville and Anniston, Ala.; Tallulah Falls 
and Bremen, Ga. 

Genus Lithobius Leach. 

Several of the species listed under this genus below conform 
to Monotarsobius as defined by Verhoeff in having the anterior 
tarsi (those of the first thirteen pairs of legs) undivided whereas 
those of species belonging to Lithobius proper have the tarsi 
all biarticulate. However, this character seems variable to 
such an extent that it is difficult to place some species uj^on 
this basis; hence, it seems best not to maintain it until some 
correlated characters, if such exist, shall be worked out. Those 
species which have the anterior tarsi clearly undivided are 
specially indicated below. ' 

6. Lithobius coecus Bollman. 

ISSS. Lithobius coccus, Bollman, Ann. X. Y. Acad. Sci.. p. 111. 

Locality. — Saluda, N. C. The only other known locality 
for this species is that at which the types were collected, Beaver 
Creek, Tenn. 

7. Lithobius tuobukus sp. nov. 

Light brown to yellow, the posterior segments often darker; head 
concolorous with body or often a darker, brown; antennae 
light brown proximally, paler distad; legs light brown, the posterior 
pairs yellow, especially bright distad. 

Antennae short or moderate; articles 2.")-2i); all except the first few 
moderate or short in length. 

Prostemal teeth mostly 5 +.5 or G+Ci, small, even. 

Ocelli compactly arranged in an oblong patch in three series; thus 
1+4, .5, 4, a total' of 14. 

None of the dorsal plates with the posterior angles produced. 

Coxal pores rather small, round, 3, 4, 4, 3. 

Last two pairs of coxae armed laterally', the last four pairs dorsally. 

Spines of the first legs 2, 2, 1; of penult 1, 3, 3, 2, the claw .single; 
of the anal 1, 3, 3, 2, the claw also single. 

1911] Lithobiomorpha of Southeastern States 37 

In the male the anal legs are moderately crassate, especially the 

third and fourth joints; the fourth joint is flattened or somewhat 

excavated dorso-mesally and is often produced at distal end into a lobe 

: extending mesad and bearing at its apex a spine directed caudad, but 

in many this lobe is absent. (See PL 3, fig. 7). 

Claw of the gonopods in female entire; spines 2+2. 

Length 9.5-12 mm. 

Localities. — Brown's Summit, N. C; Natural Bridge, Cha- 
tham, and Lynchburg, Va. ; White Suplhur, W. Va. ; Hot Springs, 
Linville Falls, Asheville, N. C; Greenville, S. C; Russellville, 
Johnson City and Unaka Springs, Tenn. ; Lexington, Ky. 

Very close to species 12 but the penult legs with only one 
claw instead of with three. Mr. Bollman mentions no modi- 
fication in the anal legs of the male of L. proridens nor does 
the specimen listed below under this species present such. 
The only course open at present, therefore, seems to be to sep- 
arate the present species from proridens and possibly to assume, 
jtidging from localities given for proridens, that Bollman has 
included the two species under one name. 

8. Lithobius watovius sp. nov. 

General color yellow; head, prehensorial feet and posterior segments 
darker, orange; antennae and legs yellow. 

Antennae short, composed of twenty articles which, excepting the 
first two and the ultimate, are moderately short. 

Prosternal teeth 2+2 or 3+3, the outer tooth on (?ach side weak 
or obsolete. 

Ocelli about four, arranged in one or two series; thtis 1+2. 1. 

Angles of none of the dorsal plates produced. 

Tarsi of the first thirteen pairs of legs undivided, those of the last 
two pairs biarticulate as usual (Monotarsobius). 

Coxal pores 1 (2), 2, 2, 2. 

Ultimate pair of coxae laterally anned, the last three pairs dorsally 

Spines of the first legs 0, 2, 1-1, 2, 1, (2) ; of penult 1, 3, 3, 0, without 
supplementary claw; of anal 1, 3, 1, 0, also without supplementary 

Anal and penult legs in the male unifomily crassate. 

Length 6.7 mm. (larger specimen). 

Locality. — Byram, Miss. Two males were taken. 

9. Lithobius paitius sp. nov. 

Dorsum pale brown; head and posterior segments darker, dark 
orange; legs greyish, except the caudal pairs which are bright yellow, 
with the brush of hairs on anal legs of male red proximally and yellow 
distad; antennae grey to dull yellow; venter pale grey to greyish yellow. 

38 Annals Entomological Society of America [Vol. IV, 

Antennae short, consisting mostly of twenty-four articles which are 
short excepting the first two and the ultimate. 

Ocelli small, in a small patch; in number about se\-en, arranged 
thus, 1+3, 3. 

Prostemal teeth 2 + 2. 

Angles of none of the dorsal plates produced. 

Coxal pores 2, 4, 4, 3, small. 

Last pair of coxae laterally armed, last two pairs dorsally armed. 

Tarsi of the first thirteen pairs of legs undivided (Monotatsobius). 

Spines of the first legs 0, 1, 1 ; of the penult 1, 3, 2, 1, without supple- 
mentary claw; of anal 1, 3, 2, 0, also without supplementary claw. 

In the anal legs of the male the fourth joint is strongly swollen and 
pro\-ided at jjroximal end with a lobe on dorso-mcsal surface from the 
posterior surface of which springs a dense brush of very long hairs which 
projects beyond the caudal end of the joint. (See PI. 3, fig. (i). 

In the female the claw of the gonopods is bluntly tripartite, the 
lateral lobes being not much lower than the middle one; basal s]iines 
as usual, 2+2. 

Length R.5-7 mm. 

Locality. — Catawba, N. C; Unaka Springs, Tenn. 

10. Lithobius watsuitus sp. nov. 

Dorsum light brown; head much darker, reddish brown or chestnut; 
prostemum colored like head, but its feet pale distad; antennae dark 
brown, paler distad; venter with the anterior plates commonly with 
purplish tinge; most legs light yellowish brown, but the caudal pairs 
darker, brown, excepting tarsi which are light. 

Antennae short; composed of thirty to thirty-two articles \\-hich, 
beyond the third arc short and compactly united. 

Ocelli about eight, arranged in two .series; thus 1+4, 3. 

Prostemal teeth, 2+2. 

Angles of none of the dorsal ]3lates produced. 

Coxal pores small and round, 2, 3, 3, 3-3, 3, 3, 3. 

Ultimate coxae laterally as well as dorsally armed. 

Spines of the first legs 1,2, 1 ; of penult 1, 3, 3, 1, one supplementary 
claw present; of anal 1, 3, 2, 0, the claw single. 

Anal and penult legs in the male crassate, especially so the fovu"th 
joint which is somewhat flattened dorso-ventrally and is longitudinall\- 
weakly furrowed. 

Length 7.5-9 mm. 

Localities. — Atlanta, Ga.; Natural Bridge, Va. The speci- 
men from Virginia differs in having the spines of the anal legs 
1, 3, 2, 1, instead of 1, 3, 2, 0. 

1911] Lithobiomorpha of Southeastern States 39 

11. Lithobius bilabiatus Wood. 

1867. Lithobius bilabiatus, Wood, Proc. Phil. Acad. Sci., p. 130. 
1887. Lithobius tuber, Bollman, Proc. U. S. N. M., p. 256. 

Localities. — Canton and Byram, Miss. This species is 
found in the states along the Mississippi river from the Gulf 
to Wisconsin and Minnesota. It seems to be most abundant 
in Illinois and Iowa. 

12. Lithobius proridens Bollman. 

1SS7. Lithobius proridens, Bollman, American Naturalist, p. 81. 

1887. Lithobius proridens, Bollman, Proc. U. S. N. M., p. 258. 

Locality. — Watervalley, Miss. One male agreeing fully 
with the original description. Previously reported from Indi- 
ana (type locality), Wa.shington, D. C; Arkansas, and Ten- 
nessee (Knoxville, Mossy Creek). 

V.i. Lithobius branneri Bollman. 

1888. Lithobius branneri, BoHman, Ann. \. Y. .■Xcad. Sci., p. 107. 
1888. Lithobius branneri, Bollman, Proc. U. S. N. M.. p. Ill, 112, 


Localities. — Brookhaven, Miss. (var. a); Maplesville, Ala.; 
Atlanta, Ga. (var. b); Catawba and Brown's Summit, N. C; 
Asheville, N. C. (var. c) ; Russellville and Unaka Springs, 
Tenn. (author). Also Knoxville, Beaver Creek, and Mossy 
Creek, Tenn. (J. C. and C. B. Branner, seq. Bollman). 

Several closely related varieties are represented in the 
material here referred to this species. The incompleteness of 
the original description must make it doubtful which variety 
is typical until the types are re-studied. The species has the 
anterior tarsi undivided (Monotarsobius). 

14. Lithobius lundii Mcinert. 

1886. Lithobius kindii, Meinert, Mvr. Mus. Haun., HI p. 111. 

1887. Lithobius lundii, Bollman, Proc. U. S. N. M., p. 111. 

Localities. — Lula and Tallulah Falls, Ga. ; Taylor's, S. C. ; 
Asheville and Hot Springs, N. C. ; Johnson City and Unaka 
Springs (and also Beaver and Mossy Creeks, seq. Bollman), 
Tenn. ; Natural Bridge. Va. 

This species, originally described from New York State, 
ranges into the southern states along the uplands. 

40 Annals Entomological Society of America^ [Vol. IV, 

15. Lithobius exiguus Meinert, var. 

ISSti. Lithobius exiguus, Meinert, Myr. Mus. Haun, III, p. 110 (11) . 
1911. Lithobius exiguus, Chamberlin, Canad. Ent. 

Localities. — Longbeach, Brookhaven, Canton, Jackson, and 
Holly Springs, Miss.; Selma (var. b), Thomasville, Morgan, and 
Birmingham, Ala.; Jackson, Tenn.; Lexington, Ky. ; Lynch- 
burg, Va. 

A widespread species occurring commonly under leaves and 
sticks and among stones along streams in Wisconsin, Illinois, 
Iowa, and neighboring states as well as throughout the region 
covered in the present paper. 

16. Lithobius elattus Bollman. 

1888. Lithobius elattus, Bollman, Proc. U. vS. N. M., XI, p. 348. 

Localities. — Johnson City and Russellville, Tenn.; Chatham, 
Lynchburg, Natural Bridge, and Balcony Falls, Va. (also 
Marksville, Va., and Washington, D. C, seq. Bollman); White 
Sulphur, W. Va. 

The specimens listed here differ somewhat from those 
described by Bollman in one or two particulars but probably 
represent the same species. 

17. Lithobius aureus McNeil. 

1887. Lithobius aureus, McNeil, Proc. U. S. N. M., p. 327. 

Locality. — Pensacola, Fla. (seq. McNeil). The two speci- 
mens upon which this species was based lack the anal legs. 
As a result it is difficult to identify the species from the pub- 
lished description. 

IS. Lithobius pinguis Bollman. 

1888. Litliubius pinguis, Bollman, Kntom. Americana, IV, p. 7. 

Localities. — Hudsonville, Miss.; (Little Rock, Ark., the 
type locality, seq. Bollman). 

Because of the incompleteness of the original description 
and the fewness of the specimens upon which based the refer- 
ence of the specimens in hand to this species is provisional. 
It is possible that the following species may have to be merged 
with the present one; but in view of the important differences 
between the specimens of L. euthus and Mr. Bollman's des- 
cription, this union at 'present seems impossible. 

19. Lithobius euthus Chamberlin. 

1904. Lithobius euthus, Chamberlin, Proc. Acad. Sci. Phil., p. G.i2. 

Localities. — Byram, Canton, and Gulfport, Miss. 

191 IJ Lithobiomorpha of Sonthcustern States 41 

20. Lithobius cantabrigensis Meinert. 

1885. Lilliobius cantabrigensis, Mriiicrt, Proc. Aiikt. l^liil. Soc, 

XXI, p. 177. 
1888. Lithobius cantabrigensis, Bollman, Proc, U. S. N. M., XI, 

p. 342. 

Localities. — Greenville, S. C; vSaluda, N. C. ; Balcony 
Falls, Va.; Russellville, Tcnn.; (also Beaver and Mossy Creeks, 
Tenn., seq. Bollman). 

Described originally from Mass., the only other recorded 

'The species seems to have a strong tendency toward the 
formation of local varieties. 

21. Lithobius cantabrigensis var. suitus, var. nov. 

Dorsuin brown; tlic head and posterior .segments darker, reddish; 
antennae pale distally; legs ])alc brown, the ])osterior pairs darker hut 
with their distal joints distinctly lighter. 

Antennae moderate, com]X)sed of from twenty-nine to thirty-two 

Ocelli about eight or nine arranged in two or three series; thus, 
1+4, 4, or 1+3, 3, 1. 

Prostcrnal teeth 2 + 2. 

Angles of the eleventh and thirteenth dorsal plates produced or in some nearly straight. 

Coxal pores round, 3, 4, 4, 3. 

Posterior coxae unarmed. 

Spines of the first legs 0, 0, 1-0, 1,1; of the penult 1,3,2,1 1 , 3, 3, 1 , 
with two claws; of the anal, 1, 3, 2, 0, with two claws. 

Gonopods in the female with the claw tripartite or almost bipartite 
in some through reduction of one lateral lobe. 

Length 7-9 mm. 

Localities. — Hot Springs, N. C; Birmingham, Ala. 
The anterior tarsi in part seem consolidated or undivided, 
but the form scarcely would conform to Monotarsobius. 

22. Lithobius cantabrigensis var. zinus var. nov. 

Color brown; head and posterior segments darker, not reddish; 
antennae dark, paler distally. 

Antennae composed mostly of from twenty-eight to thirty-one 
articles, more rarely thirty five or even thirty-seven. 

Ocelli eleven to sixteen, arranged in three or four series; thus, 
1+4, 4, 4, 3-1+3, 4, 3. 

Ultimate coxae laterally armed. 

Spines of the first legs 1, 1, 1-1, 2, 1 ; of penult legs, 1,3,3, 1-1, 3,3,2, 
with two claws; of the anal legs 1, 3, 2, 0-1, 3, 2, 1, likewise with two 

42 Annals Entomological Society of America [Vol. IV,. 

In the male the anal and penult legs moderately crassate, the fourth 
joint in the anal ones larger and somewhat complanate dorsally. 
Length S-10 mm. 

Localities. — Talapoosa and Bremen, Ga. ; Anniston, Ala. 
(variant); Brown's Summit, N. C; Chatham, Natural Bridge, 
and Lynchburg, Va. 

24. Lithobius atkinsoni Bollman. 

1.S87. Lithobius atkinsoni, Bollman, Proc. U. S. N. M., X p. G25. 
1888. Lithobius atkinsoni, Bollman, Proc. U. S. N. M., XI, p. 349. 

Bremen, Atlanta, Litla, and Tallulah Falls, (also Macon, 
seq. Bollman), Ga. ; Taylor's, Greenville and Seneca, S. C; 
Saluda and Hot Springs (also Balsam seq. Bollman), N. C. 

The localities here indicated are all those thus far recorded 
for this species. 

24. Lithobius naiwatus sp. nov. 

Brown; the head and posterior segments darker; antennae pale dis- 
tad; legs yellowish to whitish brown; the anal and sometimes also the 
penult legs dark purplish brown or purplish black, with the distal 
joints pale. 

Antennae of moderate length or short; composed of thirty-two to 
thirty-five articles of which most of the first ten are of medium length 
and those more distad short. 

Ocelli about thirteen, com])actlv arranged in three straight series; 
thus, l+,5, 4, 3. 

Prosternal teeth 2+2. 

Posterior angles of the ninth, ele\-cnth and thirteenth dorsal plates 

Coxal pores small, round, 4, .3, 5, 4. 

Last two pairs of coxae arnied laterally, last three pairs armed 

Spines of first legs, 1, 2, 1-2, 2, 1 ; of the penult legs 1,3, 2., 1-1 , 3, 2, 0, 
with two claws; of the anal legs 1, 3, 3, 2, with a .single claw. 

Anal and penult legs in male moderately crassate, without special 
lobes or processes. 

Claw of the gonopods in the female tripartite; basal spines 2+2, 
conical, the inner smaller. 

Length 11-13 mm. 

Localities. — Saluda, Catawba, and Linnville Falls, N. C; 
Landrum, S. C; Tallulah Falls, Ga. ; Unaka Springs, Tenn. ; 
Lexington, Ky. (var.) 

25. Lithobius forficatus (Linneus). 

1758. Scolopendra forficata, Linneaus, Syst. Nat., I, p. 638. 

1815. Lithobius forficatus, Leach, Tr. Linn. Soc, XL 

1821. nee Lithobius spinipes Say, Journ. Acad. Sci. Phil., II, p. lOS. 

1845. Lithobius americanus, Newport, Tr. Linn. Soc, XIX, p. 365. 

1911] Lithobiomorpha of Southeastern States 43 

Localities. — Greenville, S. C. ; Asheville and Hot Springs, 
N. C; Lynchburg and Balcony Falls, Va.; White Sulphur, W. 
Va.; Fulton and Lexington, Ky. 

The range of this species, so abundant in the north, is 
carried southward into our present territory by the mountain 
ranges. It has not previously been reported from the Caro- 
linas or other points so far south in this district. 

2(i. Lithobius celer Bollman. 

188S. Litliobius celer, Bollman, Entom. Amer., IV, p. 7. 
1909. nee. Lithobius eeler, Chamhcrlin, Ann. Ent. See. America, 
p. 190, 

Locality. — Fulton, Ky. 

27. Lithobius oedipes Bollman. 

188S. Lithobius oedipes, Bollman, Entom. Amer., IV, p. 8. 

Locality. — Mississippi. 

28. Lithobius manegitus sp. nov. 

Dorsum dark brown; head darker, nearly mahogany, the antennae 
similar proximally but becoming paler or rufous distad legs brown above, 
mostly paler vcntrally, and the posterior pairs mostly ]3ale distad. 

Antennae moderate; composed of twenty articles which decrease 
in length from the second distad to the penultimate. 

Ocelli in a patch situated apparently closer than usual to the base 
of antenna; in number about nine, arranged in three series; thus 1+3, 
3, 2. 

Prosternal teeth 2 + 2; a characteristic stout spine uniformly present 
on each side eetad of the outer tooth. 

Angles of the ninth, eleventh and thirteenth dorsal plates produced. 

Coxal pores round, in number .'i, 5, 5, 4. 

Last two pairs of coxae anned laterally; oiily the ultimate coxae 
anned dorsally. 

Spines of the first legs 1, 2, 1-2, 2, 1; of the penult 1, 3, 3, 2, with 
two claws; of the anal 1, 3, 3, 1, provided also with two claws. 

In the male the anal legs are crassate and the tibia or fifth joint is 
conspicuously furrowed lengthwise dorsally toward the exterior side, 
the furrow being fringed on each side by a dense growth of hair, the 
hair longest at posterior end (See PI. 4, fig. 7). The penult legs 
more crassate thaii the anal, the fourth and fifth joints most enlarged; 
the fifth joint or tibia furrowed from end to end along the meso-ventral 
surface and excavated on this surface at the distal end where there is a 
process bearing a conspicuous brush of hairs which projects mesad 
(See PI. 4, figs. 4, o and (J.) 

The gonopods of female with the claw entire; basal spines 2 + 2,. 
conical, the inner the smaller. 

Length 15-17 mm. 

44 Annals Entomological Society of America [Vol. IV, 

Localities. — Hot Springs, Catawba, Saluda and Linville 
Falls, N. C; Johnson City, Unaka Springs and Altapass, Tenn. 

Very close to L. oedipes Boll., but differing markedly in 
characters of the anal and penult legs of the male. 

29. Lithobius tabius sp. nov. 

Brown; head and commonly also the first dorsal plate darker, 
chestnut; antennae dark, pale distad; legs a much paler brown, the pos- 
terior pairs darker with the distal joints pale. 

Antennae short ; composed of about thirty-three articles. 

Ocelli sixteen, arranged in four scries; thus, l-|-4, 4, 4, ,3. 

Prosternal teeth 2 + 2. 

Angles of the ninth, eleventh and thirteenth dorsal ])lates produced, 
those of the sixth and .seventh exci.scd or obliquely truncate. 

Coxal pores round, 4, 4, 4, 3. 

Last two pairs of coxae laterally armed, last four i^airs anncd dor- 

Spines of first legs 1, 2, 1; of the penult 1, 3, 2, 1, with two claws; 
of the anal 1, 3, 3, 2, with two claws. 

Claws of the gonopods in female tripartite. 

Length 10.5 mm.; width at tenth dorsal plate 1..5 mm. 

Locality. — Johnson City, Tenn. 

In many points very similar to arienus, carolinae, etc., but 
the angles of the sixth and seventh dorsal plates not at all 

30. Lithobius simitus sp. nov. 

Brown; the head and posterior plates reddish, the fonner paler 
cephalad of the frontal suture; antennae dark brown, pale distally; 
legs whitish brown, the tarsi clear yellow, the posterior pairs darker, 
the anal pair yellow distad of the femur; venter light brown, darker 
caudad, the anterior plates with purplish tinge. 

Antennae short, composed of twenty-seven to thirty-two short 

Ocelli arranged in two series, about eight in number; thus, l-|-4, 3. 

Prosternal teeth 2-|-2. 

Angles of the ninth, eleventh and thirteenth dorsal plates produced. 

Coxal pores small, 2, 3, 3, 3. 

Last two pairs of coxae anned laterally, last three pairs armed 

Tarsi of the first thirteen pairs of legs imperfectly divided in part 
but suture mostly plainly evident. 

Spines of the first legs 0, 0, 0-0, 0, 1; of the penult 1, 3, 2, 0, with 
two claws; of the anal 1, 3, 2, (), a supplementary claw likewise present. 

Claw of the female gonopods bipartite; basal spines 2+2, conical. 

Length 7-7.5 mm. 

Locality. — Grenada, Miss. Two female specimens. 

1911] Litbobiunwi'pha of Soiilheasteni States 45 

31. Lithobius transmarinus Koch. 

lsii2. Litlidbius iransmarinus, Koch, Die Myriopodcongattung Litho- 

Ijius. p. 31. 

. Lithobius mordax, Koch, ibid. p. .34. 

1872. Lithobius mordax, Meinert, Myr. Mus. Haun., II p. 294. 
1875. Lithobius transmarinus, Stuxberg, Ofvers. af K. Vet. Akad. 

Forh. no. 3, pp. 2G and 32. 

. Lithobius morclax, Stuxberg, ibid., pp. 27 and 32. 

18S7. Lithobius mordax, Bolhnan, Proc. U. S. N. M., p. 263, etc. 

. Lithobius transmarinus. Bollman, ibid., p. 626, etc. 

1893. Lithobius spinipcs, Bolhnan, (of Say??), Bull. U. S. N. M.. 

46, p. 146. 
1896. Lithobius mordax var. Louisianac, Brolemann, Ann' Soc. 

Ent. de France, p. 48. 
. Lithobius transmarinus var. pennatus, Brolemann, ibid., p. 48. 

Localities. — Brookhaven, Fernwood, Holly Springs, Byram, 
Canton, Biloxi, Ocean Springs, and Longbeach, Miss.; New 
Orleans, La.; Jackson, Mobile, and Salem, Ala. 

Apparently Koch based his description of transmarinus upon 
a female while his description of mordax is clearly that of the 
male. The differences pointed out between the anal legs of 
these two forms are essentially secondary sexual characters as 
found in male and female of the present species, although the 
longitudinal furrows on the mesal surface of the sixth and 
seventh joints in the female vary in development and may be 
deeply impressed in some, in others evident upon one or the 
other of the joints alone, or may be quite absent as seems to 
be more commonly the case in the males. vSimilarly Brolemann 
(Ann. Ent. Soc. Fr., 1S96, pp. 4S-49) in arguing for the dis- 
tinctness of transmarinus and mordax relies almost wholly 
upon secondary characters and does not inform us as to whether 
he is speaking of male or female. He says: "Pour ce qui est 
des deux especes de Koch, il me semble qu'il ne pent y avoir de 
confusion, puisque le 4e article des pattes anales des mordax 
est tres court, tres renfle, parcouru en dessus par un profond 
et large sillon, ce qui n'est nullement le cas chezle transmarinus." 
But this is true only of males while the females conform to 
Brolemann's transmarinus, which must accordingly be regarded 
as the same species. 

32. Lithobius xenopus Bollman. 

1S8S. Lithobius xenopus, Bollman, Proc. U. S. N. M., XI, p. 350. 

Locality. — Tallulah, Ga. (L. M. Underwood). 

33. Lithobius vorax Meinert. 

1S72. Lithobius vorax, Meinert, Myr. Mus. Haun., II, p. 292. 

1.875. Lithobius vorax, Stuxberg, Ofvers. af k. Vet. Akad. Forh. no. 

3, p. 26 and 32. 
1885. Lithobius latzeH, Meinert, Proc. Am. Phil. Soc, XXI, p. 175. 
1887. Lithobius clarus McNed, Proc. U. S. N. M., X, p. 326. 
. Lithobius tyrannus, Bollman, ibid., p. 6.36. 

46 Annals Entomological Society of America [Vol. IV, 

Localities. — Byram, Fernwood, Watervalley, Canton, H0II3- 
Springs, Grenada, Jackson, Biloxi (type locality), Longbeach 
and Ocean Springs, Miss. ; Pensacola, Fla. (clarus McNeil) ; 
Jackson and Birmingham, Ala.; Brown's Summit, N. C; 
Crandall, Marksville and Luray, Va. (latzeli, seq. Meinert and 

There is marked variation in this species in the development 
of the claws of the penult legs. There is a distinct anterior 
or third claw in the specimens from some localities (such as 
Holly Springs, Fernwood, Longbeach, Miss., and Jackson, Ala., 
etc.) which seems to be especially well developed in younger 
or smaller specimens (cf. clarus McNeil) but which is readily 
broken off and tends in older specimens to become relatively 
reduced, obsolete or absent. Because of this it would seem 
justifiable to regard clarus as having been based upon small 
specimens of this species since no other difference appears in 
the description given. Specimens from North Carolina and 
Virginia seem to show a tendency for the coxal pores to be 
round or oval rather than strongly transverse more frequently 
than in specimens from the Gulf region; but there are no con- 
stant differences in this respect and both extremes with inter- 
mediates are to be found in the more southern localities. Hence, 
no grounds in this direction appear why L. latzeli should be 
kept apart from vorax. The longitudinal sulcus which Meinert 
mentions as occurring on the ventral surface of the third and 
fourth joints of the anal legs in latzeli is present in all specimens 
of vorax. The sulcation on the mesal surface of the tarsal 
joints of anal and penult legs mentioned by Bollman as dis- 
tinctive of his tyrannus is present in most specimens of vorax. 
The articles of the antennae vary greatly in number with the 
size of the individual, from twenty-six or twenty-seven in young 
specimens fourteen or fifteen millimeters in length to above 
forty in the largest adults. The average number in medium 
size adults would seem to be about thirty-five or thirty-six. 

34. Lithobius underwoodi Bollman. 

ISSS. Lithobius underwoodi, Bollman, Proc. U. S. X. M., XI, p. 350. 

Localities. — Maplesville, Selma, Morgan, Thomasville, Jack- 
son, and Anniston, Ala.; Atlanta, Tallulah Falls, (and Macon, 
type locality, seq. Bollman), Ga. ; Landrum and Seneca, S. C. 

Especially abundant in Alabama where it seems to be the 
most common of the larger species. 

1911J LithohiomorpJia of Southeastern States 47 

35. Lithobius rex B oilman. 

1SS8. Lithobius rex, BoUman, Procl U. S. N. M., XI, p. 350. 
Locality.— Tallulah, Ga. (L. M. Underwood). 

-36. Lithobius carolinae sp. nov. 

Medium or slender. 

Dorsum brown to brownish yellow; head much darker, chestnut; 
prostemum dark brown; antennae bark brown, pale or rufous distally; 
legs light brown or yellow, the posterior pairs darker but light distally; 
venter pale. 

Antennae short; composed of thirty to thirty-five articles. 

Ocelli about ten or twelve, arranged in three series; thus, 1 +4, 3, 3. 

Prosternal teeth 2+2. 

Angles of the .sixth, seventh, ninth, eleventh and thirteenth dorsal 
plates produced. 

Coxal pores 3, 4, 4, 3, round. 

Last two pairs of coxae laterally anned, last three pairs dorsally 

Spines of first legs 0, 0, 1; of penult I, 3, 2, 1, with two claws; of 
the anal 1, 3, 2, 0, or rarely 1, 3, 2, 1, also with two claws. 

Claws of gonopods in female tripartite; basal spines 2+2. 

Length 8-9.5 mm.; width of tenth dorsal plate L2~L5 mm. 

Localities. — Asheville and Hot Springs, N. C. ; Landrum 
and Taylor's (var.), S. C; Russellville, Tenn. 

This species would seem to be related to L. juventus of 
Bollman, but the sixth dorsal plate has the posterior angles 
more or less produced and the spining of the legs is constantly 

37. Lithobius arienus sp. nov. 


Dorsum brown; head together with anterior and posterior plates 
darker, not reddish; antennae brown, darkened distad; legs pale yellow, 
the posterior ]3air darker; venter pale, the prosternum and posterior 
plates slightly darker. 

Antennae moderately long; composed of thirty-four articles. 

Ocelli compactly arranged in four longitudinal series; thus 1+4, 
4, 3, 3, a total of fifteen; ocelli of the two upper rows larger. 

Prosternal teeth 2 + 2. 

Sixth, seventh, ninth, eleventh and thirteenth dorsal plates with the 
posterior angles produced. 

Co.xal pores ad 3, 4, 4, 3, round. 

Last two pairs of coxae laterally armed, the last three pairs dorsally 

Spines of the first legs 0, 1, 1 ; of the penult 1, 3, 3, 2, armed with 
two claws; of the anal 1, 3, 2, 1, also armed with two claws. 

Length 11.5 mm.; width of tenth plate 2 mm. 

48 Annals Entomological Society of America [Vol. IV, 

Locality. — Hot Springs, N. C. 

One male. Very close to carolinae but conspicuously 
larger and more robust and differing in the spining of the legs, 
in the ocelli, in coloration, etc. 

Genus Bothropolys Wood 
But one species of this genus occurs in the United States 
east of the Rocky Mountains. — B. multidentatus. 

.38. Bothropolys multidentatus Newport. 

1845. Lithobius multidentatus, Newport, Tr. Linn. ,Soc., XIX, p. 36.5. 
1862. Bothropolys nobilis. Wood. loum. Acad. Sci. Phil. V, p. 1.5. 
1S6.5. Bothropolys multidentatus, Wood, Tr. Am. Phil. Soc., XIII, 

p. 1.52. 
1875. Lithobius multidentatus, Stuxberg, Ofvers. af k. Vet. Akad. 

1887. Lithobius multidentatus, BoUman, Proc. U. S. N. M., p. 263. 

Locahties. — Canton, Fernwood, and Byram, Miss.; Maples- 
ville and Jackson, Ala.; Catawba, N. C; Russellville, Tenn.; 
White Sulphur, W. Va. ; Chatham and Balcony Falls, Va. 

This species is widespread in the southern states though 
apparently not so common as farther north. In the section 
from Virginia to New York state, etc., it is abundant as it is 
also in corresponding latitudes farther west. 

Plate 3. 

Fig. 1. Left anal leg of Lamyctes fulyicornis from the exterior. From 
specimen 9.6 mm. in length taken at Haugen, Wisconsin. 

Fig. 2. Left anal leg of Lamyctes tiyius sp. nov. From a specimen 6.6 
mm. in length taken at Jackson, Ala. Same magnification 
as preceding. 

Fig. 3. Left anal leg of Lamyctes pinampus. From a specimen 8.6 mm. 
long taken at C'laremont, Cal. Same magnification as the 

Fig. 4. Right anal leg of Watobius anderisus. .Specimen from Thomasville, 

Fig. 5. Right penult legs of Watobius anderisus. Same specimen as preced- 

Fig. 6. Left anal leg of Lithobius paitius sp. noy., dorsal aspect. 

Fig. 7. Lett anal leg of Lithobius tuobukus sp. noy., dorsal aspect. 

Pl.\te 4. 

Fig. 1. Right leg of the fifth pair of Buethobius oabitus sp. noy., cephalic 

Fig. 2. Right leg of the fifth pair of Zygethobius pontis sp. noy. 
Fig. 3. Right leg of the fifth pair of Watobius anderisus sp. nov., cephalic 

Fig. a. Left penult leg of Lithobius manegitus sp. nov., ventral aspect. 
Fig. 5. Left penult leg of Lithobius manegitus sp. nov., dorsal aspect. 
Fig. 6. Left anal leg of Lithobius manegitus sp. nov., dorsal aspect. 
Fig. 7. Prosternum of Lithobius manegitus sp. nov., ventral aspect. 

Annals E. S. A. 

Vol. IV, Plate HI. 

X. I'. Chamberlin. 

Annals E. S. A. 

Vol. IV. Plate IV. 

K. T. CItamberliii. 


H. F. Wilson, 
Oregon Agricultural College. 

In taking up the preliminary work on what I term the sub- 
family Lacliiiinae it is very hard to ascertain the correct stand- 
ing of several of the genera in the tribe Lachnini both from a 
standpoint of literature and classification. 

Beginning with the original description of the genus Lachmcs 
the author will discuss the later genera as erected and invites 
further discussion in order that the correct generic names may 
be used in the future. 

The genus Lachnus Burmeister. Illiger is credited with 
the genus, but in reality it belongs to Burmeister and was 
published in 1835 in his Handbuch der Entomologie, p. 91. 

"3 (13) Gatt Lachnus 111. 
Aphis autor. 

Eh. Fuhler deutlich sechsgliederig, kurzer als der Leib. Das 
erste und zweite Glied kurz und dick, das dritte sehr lang, das vierte 2-3 
kurzer, das funfte etwas langer als das dritte, das sechste klein, zuges- 
]Ditzt bei einigen Artcn wic am Ende zusammen geschnurt und scheinbar 
ein eigenes Glied bildend. Flugel mit starker Randrippe. Hintcrleib 
ohne Honigrohrcn hochstcns mit zwei Hockem an deren Stelle. " 

Under this genus Burmeister gives seven species as follows: 

1. Lachnus lapidan'us, (Fab.), which appears to be an 
unrecognizable species. 

2. Lachnus fagi, (Linn.), which is now the type of the 
genus Phyllaphis Koch.' 

3. Lachnus quercus, (Linn.), which is now the type of the 
genus Stomaphis Walker. - 

4. Lachnus fasciatus, Burmeister, which Del Guercio has 
recently placed in his Genus Lachniella.'' 

5. Lachnus Punctatus, Burmeister, which up to the present 
time has not been definitely recognized (may be viminalis 
Boyer).^ (?) 

1. Koch, Die Pflanzenlause Aphiden, 1857, p. 24S. 

2. Walker, The Zoologist, 1870, Vol. 28, p. 2000. 

3. "Redia," 1900, Vol. 5, fasc. 2, pp. 173-3.59. 

4. Boyer, Ann's Ent. Soc. France, 1841, p. 184. 


52 Annals Entomological Society of America [Vol. IV, 

Then he mentions Aphis pini aut. and Aphis betidae autor to 
go in this genus but as no reference is made to any one author 
neither species can have a valid standing in this genus. 

As one of the species originally cited under the genus must 
hold for the type of that genus then must one of the four valid 
species be that type. 

Two of the four are unquestionably removed as types of 
the genera PhyUaphis and Stomaphis, thus leaving only two for 
the genus Lachnus. 

Lachniis punctatiis if found to be distinct is the only species 
which has not been definitely recognized and placed in a differ- 
ent genus by the later writers, and it is the only species left for 
the type of the genus. ° Unless this species is located the 
genus Lachnus must revert to the group containing L. fasciatus 
Burm. as a type. 

A careful study of Lachnus viminalis Boyer, Boyer's descrip- 
tion of that species, and Burmeister's description may (?) 
show that L. viminalis Boyer is identical with L. punctatiis 
Burm. In that case Lachnus will be definitely established 
with L. fasciatus as the type. If not then what is the genus 
and what species can we refer to that genus? 

On the other hand in 190S Mordilko'^ used L. viminalis 
Boyer to form a new genus Tuberolachnus. Should this species 
prove to be L. punctatus then L. fasciatus Burm. must be the 
type of the genus Lachnus Burm., as it is the only species of 
those cited by Burmeister left in that genus. Since L. fasciatus, 
according to Del Guercio at aut., is a valid species I hold that 
this species under the existing conditions must hold as the type. 

The next genus taken up in this tribe was Cinara Curtis, 
as follows: 

The genus Cinara Curtis, 
type A. pini Linn.? 
He includes A. roboris Linn. 

5. April, 1910. Entomological News. The author gave Lachnus pimclalus as 
the type of the genus Lachnus because it seemed to be the only species which was 
left for that genus, and at that time I was unaware of the fact that Mordwilko 
(Annuaire Music Zoologique de L' Academic Imperiale des Sciences, Vol. 13, 1908, 
p. 374) had used Lachnus viminalis as the type of his genus Tuberolachnus. It is 
impossible, however, with the present knowledge of the two above species to more 
than place Lachnus punctatus as a doubtful synonym of L. viminalis for Lachnus 
punctatus apparently cannot be clearly determined, and Boyer's description of 
L. viminalis is too clear to be put aside. 

6. Annuaire Musie Zoologique de L'Academie Imperials des Sciences, vol. 
13, 1908, p. 374. 

1911] Notes on Synonymy of Lachnini 53 

This genus was formed in 1835 by Curtis, section 576, Vol. 
12, of his British Entomology. 

He places two species in the genus. Aphis puii Linn.?, 
and Aphis roboris Linn. The first he gives as the type, but as 
he places a question mark after Linn., the species is not valid, 
and A. roboris Linn, which he describes in full is the type of 
the genus? The generic names erected for that species since 
that time are synonyms?' He gives the figures of the adults, 
some of the parts, and also gives a good description. 

The synonymy of this genus would then be 

Cinara Curtis 1835 
Pterochloriis Rondani 1848.' 
Dryobius Koch 1855 Loc. cit. 

Dryaphis Amyot" which Del Guercio Loc. cit. p. 262 has 
given genus rank never was a genus name until given that rank 
by Del Guercio. If we were to accept Amyot's names which 
were mononomials and in this case means "Oak Aphid" there 
would never be an end to the changing of names. The late 
workers on the Hemiptera refuse to look upon the work of 
Amyot except as a curiosity. 

The next genus to be formed in the Lachnus group was 
Stomaphis Walker loc. cit. with A. quercus Linn, as the type 
and there is no discussion necessary on this genus name as it is 
well established. 

Mordilko loc. cit. in 1908 deemed it necessary to erect 
two new genera in this group, Schizolachnus Mord. with ^4. 
tomentosus DeGeer as the type and Tnberolachnus Mord. with 
Lachnns viminalis Boyer as the type. 

In 1909 Del Guercio loc. cit. has placed both of the above 
species in the genus Lachnus regardless of the fact that neither 
were in the original genus and he removes to other genera all 
of the original included species. If it is true that L. viminalis 
Boyer and L. tomentosus DeGeer are both in the same genus 
then must Tnberolachnus be the genus name with Schizolachnus 
as a synonym and L. viminalis Boyer as the type. 

7. The question of the validity ot this genus rests upon the fact that Curtis 
did not give roborus as the type and the other species is questioned. The author 
then concludes that the genus is in question and cannot be placed as a valid genus. 

8. Esapodi afidicidi in Nuove Ann. di Sci. Nat. Bologna, 184S. 

9. Ann. Soc. Ent. France vol. 5, ser. 2, p. 481, 1847. 

54 Annals Entomological Society of America [Vol. IV, 

In 1909 five new genera were formed in this group, one of 
which must be a synonym and a second which would accord- 
ing to the reasoning of this article also be a synonym. 

The genera are Eulachniis Del Guercio (loc. cit.), the type 
of which probably should be E. Agilis (Kalt.) 

Lachniella Del Guercio (loc. cit.), the type of which is not 
set, and is, I consider, a synonym of Lachnus? 

EssigeUa Del Guercio (loc. cit.) with E. californicus (Essig) 
as the type. 

Davisia Del Guercio (loc. cit.) L. longistigma Monell as the 
type and which is a synonym of the following genus. (Nov. 
13, 1909). 

Longistigma Wilson '", type L. caryae Harris which I have 
published as synonymous with L. longistigma Monell and L. 
platimcola Riley. (Nov. 1, 1909.) 

According to the evidence shown here using Del Guercio's 
arrangement to generic characters, the correct synonymy is as 

follows : 

1. Trama Heyden 

type T. troglodytes Heyd. 

2. Stomaphis Walker 

type 5. Quercus (Linn). 

3. Picrochlorvs Rondani 

Syn. Cinarn Curtis? 
Syn. Dryobijis Koch. 
Syn. Drynphis Kirk 
type P. roboris (Linn.) 

4. Essigella Del Guercio 

t\pe E. californicus (Essig.) 
•5. Longistigma Wilson 

Syn. Davisia Del Guercio 
type L. caryae (Harris). 

6. Tnherolachnus Mord. 

? .Syn. Schizolachnus Mord. 
type T. I'iminalis (Boyer). 

7. Lachnus Burmeister. 

S5"n. Lachniella Del Guercio 
Type L. fasciatus Burm. 

8. Eulachnus Del Guercio 

type E. Agilis (Kalt). 

In the December, 1910, issue of the Anx.\ls the author 
published a paper on the genera of the subfamily Aphidinae 
and wishes here to note two corrections. 

The type of the genus Illinoia shoidd read m. liriodendri 
^lonell. The type of the genus Hyaloptents Koch should read 
A. priini Fab. instead of aurantiae Koch. 

10. Can. Ent., vol. 41, p. 385, 1907. 


By E. P. Felt, 

Albany, N. Y. 

Our iinderstanding of this group will be much clearer if 
we recall that it is an offshoot from the Mycetophilidae, the 
species of which subsist largely upon decaying vegetable matter 
or low forms of vegetable life. The family Itonidae, better 
known as the Cecidomyiidae, has attained its present large 
proportions not by reason of strength, great resistant powers 
or unusual fecundity, but through an amazing adaptability. 
We find larvae in decaying vegetable matter, in dead wood, on 
fungus, affecting all parts of a very great variety of the higher 
plants and presenting thereupon almost every conceivable 
grade in the development of the gall, living as parasites at the 
expense of very small insects or even preying upon their near 
allies. Broadly speaking, taxonomic studies in this family 
show at least a moderately close relationship between speciali- 
zation in structure and divergence in the food habits from those 
of ancestral forms. 

We have no firsthand knowledge of the food habits of the 
tribe Lestremiinariae, though there is every reason to believe 
that there is substantial agreement in this regard with Euro- 
pean species, which have been reared from decaying 
vegetable matter. 

We can supply a little definite information respecting the 
food habits of the tribe Campylomyzariae, since representatives 
of several genera have been reared. Mycophila fungicola, an 
undescribed species, referable to a new genus was reared from 
fungus, while Monardia lignivora Felt was obtained in consid- 
erable numbers from the fungous-affected heartwood of white 
pine. Cordylomyia coprophila is an undescribed species refer- 
able to a genus which will be erected shortly. It was reared 
from manure. These few records show that this compara- 
tively generalized tribe subsists upon fungi, fungous-affected wood 
and certain forms of vegetable matter. These food habits agree 
in general with those of European species, and further observa- 
tions will doubtless show that members of the tribe as a whole, 
depend for nourishment on the lower plants or upon the tissues 


56 Annals Entomological Society oj America [Vol. I\', 

of the higher plants after invasion b}- fungi or the commence- 
rnent of decay. 

The subfamily Heteropezinae comprises a number of remark- 
able, and taxonomically speaking, ancient forms. The species 
live largely, if not exclusively, in ligneous tissues in the incipient 
stages of decay. Miastor larvae, presumably those of M. amer- 
icana Felt were found in numbers in the moist, partially decayed 
inner bark and sapwood of chestnut. The majority of the 
European species studied, develop under practically similar 

Our title implies a limitation to gall-making forms. This 
is true of the vast majority of the members of this family, 
though not applicable to the two subfamilies just discussed 
or to the lowest tribe of the Itonidinae now under considera- 
tion. The members of this tribe, the Epidosariae, distinctly 
allied with the more generalized forms in this family by the 
presence of a well developed crossvein and yet exhibiting a 
connection with the higher forms because of the universal 
presence of highly specialized circumfili, do not produce galls 
but live in dead, frequently dried, woody tissues. The detection 
of the larvae is consequently difficult and, as a result, rearings 
have been comparatively few. Winnertzia pinicorticis Felt 
was obtained by Mr. Pergande from the bark of Pinus inops. 
The genus Colpodia, with its remarkably long, narrow wings, 
probably lives in dead wood, a habit known to be true of 
Asynapta saliciperda Felt which was reared from old Rhab- 
dophaga batatas O. S. galls on willow. 

The most generalized of the true gall-making forms are 
probably found in the tribe Dasyneuriariae, the genus Rhab- 
dophaga Westw. being the less specialized of this group. A 
study of this genus shows at once a marked partiality to Salix, 
a genus placed rather low in the series of flowering plants, and 
the production thereupon of a number of comparatively simple 
deformities such as bud and subcortical galls. It is pre-emi- 
nently a genus of the willow. Dasyneura Rond. comes next. 
An examination of the records shows that a large proportion of 
the species live in comparatively simple leaf and bud galls on 
A'arious genera of the higher flowering plants, an interesting 
exception being the remarkable D. flavotibialis Felt which 
was reared from decaying wood, while D. rhois Coq. was 
obtained from a root gall on Sumac. The peculiar Lasiopteryx 

lull] Food Habits of American Gall Midges 57 

coryli Felt was reared from leaf folds on hazel, Corjdus virginica. 
An undescribed species of Cystiphora Kieff. was reared from a 
very inconspicuous swelling on Viburnum leaves. A departure 
from the normal food habit in this tribe is seen in Coccidomyia 
pennsylvanica, an undescribed species belonging to a new genus 
and reared from Lecanium scales. 

This series of generalized gall-making forms is continued in 
the tribe Oligotrophiariae, separated from the preceding only 
by the simple claws. The more generalized genus, Phytophaga 
Rond. exhibits a connection with the preceding tribe in the 
possession by P. destructor Say., of claws with rudimentary 
teeth. As is well known, it hardly makes a gall, depending for 
protection upon the leaf sheath. P. ulmi Beutm. occurs in 
the buds of elm, P. violicola Coq. curls the leaves of violet, 
while several species of this genus typified by P. rigidae O. S. 
live upon vSalix, making galls similar to those produced by spe- 
cies of Rhabdophaga. Janetiella asplenifolia Felt was reared 
from a fleshy fold on the midvein of sweet fern, while J. brevi- 
cauda Felt was obtained from the typical gall of Lasioptera 
vitis O. S. on grape. The genus Oligotrophus Latr. is repre- 
sented by the European O. betulae Winn., which affects the seeds 
■of birch, while O. salicifolius, an undescribed species produces 
a flatfish, ovoid gall on Salix leaves. The genus Rhoi:)alomyia 
Rubs, contains a large number of species and exhibits a marked 
partiality for Solidago, producing upon various species of this 
plant genus a considerable number of flower and bud galls, the 
large rosette deformities of apical buds being characteristic. A 
few species of this genus also occur upon the allied aster and 
Artemisia. The larger species of Sackenomyia Felt are re- 
stricted to Salix, while one small species at least, has been reared 
from Viburnum. Walshomyia Felt is found v\ the fruit of 

The tribe Lasiopterariae exhibits a high degree of specializa- 
tion in venation at least, and we find in this group a marked 
restriction in food habits. The genera Lasioptera Meign. and 
Neolasioptera Felt live almost exclusively in subcortical stem 
galls, a large proportion of the species occurring upon Solidago, 
though a considerable variety of other plants are subject to 
attack. Asteromyia Felt, like the two preceding genera, 
exhibits a marked preference for Solidago, though a number of 
species occur upon aster. It is noteworthy that a large major- 

58 Annals Entomological Society of America [Vol. IV, 

ity of the galls produced by this genus are of the apparently 
fungous-affected blister type. The highly specialized Clinor- 
hyncha Loew is represented in America by several species 
probably restricted to the florets of Yarrow, Thoroughwort 
and presumably Chrysanthemum. The peculiar Campto- 
neuromyia adhesa Felt has been reared from oval, adherent 
galls between Solidago leaves, while C. rubifolia Felt was 
obtained from a marginal leaf roll on blackberry. 

The tribe Asphondyliariae is a rather highly specialized 
group, the species living mostly in buds. This is particularly 
true of Asphondylia H. Lw., a genus practically confined tO' 
buds and apparently not closely restricted in food habits, 
since different species have been reared from a considerable 
variety of plants. Schizomyia Kieff. is allied to the preceding 
genus and the several species reared were obtained from buds; 
such as S. coryloides Walsh and Riley from an apical leaf bud 
gall on grape, and S. pomum Walsh and Riley from a nutlike 
polythalamous grape gall, evidently a modified bud. S. rivinae 
Felt was reared from bud galls on Rivina. Cincticornia Felt 
appears to be restricted to leaf galls on Quercus, the largest and 
perhaps most characteristic being that produced by C. pilulae 
Walsh. A series of rearings have resulted in obtaining a number 
of species, all from various leaf galls on this plant genus. 

The tribe Itonidinariae comprises a large assemblage of 
highly specialized forms, easily divided by the circumfili into 
two groups, namely the bifili and trifili. The former is repre- 
sented by Endaphis Kieff. first recorded as an .endoparasite 
on Aphididae and reared by us from mite infested foliage. 
Contarinia Rond. also belongs in this subtribe and, as is well 
known, displays a marked preference for bud and fruit structures, 

C. johnsoni SHng., C. virginianeae Felt, C. rumicis H. Lw., C. sor- 
ghicola Coq. and C. pyrivora Riley, all being representative in 
food habit. Thecodiplosis Kieff. is closely allied to the preceding 
and is represented in America by T. quercifolia Felt reared from 
oak, T. ananassi Riley reared from a twig gall on Taxodium, 
and T. liriodenri O. S., inhabiting a blister gall on tuHp leaves. 
Dentifibula Felt, also in this subtribe, has at least one species, 

D. cocci Felt, which is zoophagous. 

1911] Food Habits of American Gall Midges 59 

The subtribe trifili comprises the remainder of the genera 
in the family. The genus Bremia Rond., representated by sev- 
eral American species, is probably phytophagous. Aphidoletes 
Kieff. contains several American species, a few of which at least 
are known to prey upon Aphididae. It is possible that our 
American species of Lobodiplosis Felt, Coquillettomyia Felt, 
and Karschomyia Felt have habits similar to those of 
the allied Mycodiplosis Rubs., the majority of the species of 
which appear to subsist upon fungi, though one, M. acarivora 
Felt preys upon Tetranychus. Youngomyia Felt displays a 
preference for the buds of various plants. Species of Clino- 
diplosis Kieff. have been reared from leaf galls on scrub oak. 
Spiraea, Carya, and from roots of Cattleya. It is probable that 
the species occurring on hickory leaves is an inquiline. The 
genus Caryomyia Felt comprises a number of homogeneous 
forms producing a considerable variety of galls on hickory 
leaves. We have yet to obtain undoubted evidence that mem- 
bers of this genus live upon any other plant. Prodiplosis 
floricola Felt has been reared from enlarged blossoms of spiraea 
and clematis. Arthrocnoda.x Rubs, is represented by several 
American forms, A. apifila Felt occurring in bee hives and prob- 
ably subsisting upon organic debris, though subsequent inves- 
tigations may show it to be predaceous. Hormomyia H. Lw. 
comprises a number of large forms usually found in the vicinity 
of swamps and presumably living mostly on sedges or allied vege- 
tation. Four species, hardly typical of the genus, namely H. 
crataegifolia Felt, H. canadensis Felt, H. clarkei Felt and H. ver- 
ruca Walsh have been reared from leaf galls respectively, on 
Crataegus, Amelanchier, Spiraea and Salix. The European 
Monarthropalpus buxi Lab., producing an oval swelling upon the 
leaves of Box has been recently detected in this country. Giar- 
domyia menthae Felt was reared from a pustule-like gall in the 
axil of the leaf of Mentha canadensis. Lestodiplosis Kieff. 
is represented by a large series of mostly spotted-winged 
midges which have been reared from a considerable variety of 
plants. The larvae of some at least, are known to be zoopha- 
gous and it is probable that most of the reared American forms 
prey upon the larvae of gall-making midges. The genus 
Itonida Meign,. better known as Cecidomyia Meign., comprises 
a large number of forms inhabiting for the most part, flower, 
bud and leaf galls on the higher flowering plants, though L 

()0 Annals Entomological Society of America [Vol. IV', 

resinicola O. S. and I. resinicoloides Wlms. occur in exuded pitch 
masses on pine, while I. tritici Kirby is well known as a species 
of prime economic importance. 

A study of our records from a botanical aspect reveals 
several facts of interest. We note first that American gall 
midges live at the expense of some 177 plant genera belonging 
to 66 plant families. They afford support to some bZS 
species of gall midges representing 44 genera. These forms 
are known to inhabit 44 fruit (botanically speaking), 146 bud, 
218 leaf, 130 stem, and 4 root galls. The paucity of root galls 
must be attributed in a measure to the difficulty of finding them. 
In addition to the above some five species were reared from 
unknown plants and eleven zoophagous species belonging to 
three genera, making a total of 47 insect genera comprising 
some 5.54 species, 441 of these having been reared from either 
plants or animals. Reference to our records shows that the 
Compositae supports a very large fauna, 22 of its genera afford- 
ing sustenance to 118 species of gall midges belonging to some 
15 genera. The majority of these midges, 55 species occur in 
bud, 32 in leaf, 30 in stem, while 5 inhabit fruit galls. The 
Salicaceae, represented only b}^ Salix and Populus, supports 
some 59 species of gall midges referable to 15 genera, by far the 
greater number occurring upon Salix. As in the Compositae, 
a large proportion, 21 species occur in bud, 15 in leaf 
and 21 in stem galls, only 1 living at the expense of 
the fruit. The Rosaceae appca^s to be the next plant 
family favored by gall midges, 10 genera being sub- 
ject to attack by 43 species of midges, assignable to 14 
genera, 3 species inhabiting fruit, 12 bud, 25 leaf and 
only 3, stem galls. The Gramineae, despite its numerous genera 
and wide distribution has but 18 genera at present known to 
support some 25 species of midges representing 12 insect genera, 
8 species occurring in fruit and 17 in stem galls. This is prob- 
ably only a small proportion of the forms occurring upon grasses. 
Our record for the Cyperaceae is even more unsatisfactory, only 
one species, presumably inhabiting a stem gall being known. 
The paucity of records in both of these families is probably 
due to the difificulty of finding the galls. There is a close paral- 
lelism between the Juglandaceae and the Fagaceae, two genera 
in each being affected by gall midges. Juglans and Castanea 
are known to be infested by one and three species respectively, 

1911] Food Habits of American Gall Midges (il 

while Carya and Quercus arc suVjject to attack by 25 and 21 
species, the former affording support to representatives of 
5 and the latter to species belonging to 7 genera. These two 
trees are likewise comparable in that each supports but one 
species in the fruit, while by far the greater majority of the 
midges, namely 23 and 18 respectively, produce leaf galls. 
The large family Legumin9sae has 13 genera which support 
some 5 genera of gall midges referable to 20 species, 3 living in 
fruit, 3 in l)ud, 6 in leaf and 8 in stem galls. Only 6 genera in the 
Urticaceae are attacked by gall midges belonging to 4 genera 
representing 8 species, 2 living in bud, 9 in leaf and 6 in stem 
galls. In the Vitaceae, Psedra, and Vitis support some 12 
genera of gall midges representing 17 species; 4 inhabit bud, 
12 leaf and 1 root galls. By far the great majonty of the 
species, 1.5, occur upon Vitis. The large family of Labiatae 
supports some genera representing only 13 species, the 
Caprifoliaceae, 8 genera comprising 14 species, and the import- 
ant Pinaceae (i genera and 14 species. The above record, while 
dealing with a much larger number of species than we have been 
accustomed to think occurred in this family, shows that in all 
probability there are many forms yet to be discovered. 

Comparing the above data with recently summarized records* 
it will be seen that the food habits of some 420 European gall 
midges representing 43 genera are unknown. The Pinaceae 
afford sustenance to 11 species belonging to 4 genera, a condi- 
tion closely paralleled in this country. The European Grami- 
neae support some 20 gall midges representing 7 genera, a show- 
ing somewhat below what obtains in America. Conversely, 
the European records for the Cyperaceae include 4 genera and 
9 species, while in this country but one species has been reared 
from Scirpus. The European Salicaceae supports some 30 
species of midges belonging to (i genera, 5 of these occurring 
on poplar. There appear to be no species affecting the Juglan- 
daceae in Europe. There are nearly as many genera and species, 
20 and 7 respectively, occurring upon the Fagaceae in Europe 
as in America, though the distribution is different, since Fagus 
supports 5 species referable to 3 genera and Quercus has only 
14 species representing 4 genera, a marked contrast to condi- 
tions obtaining in this country. There is a pronounced differ- 

* 1909, Houard, C. Les Zoocecidies des Plantes d'Europe. 

62 Annals Entomological Society of America [Vol. IV, 

ence in the European Rosaceae, especially marked in Spiraea 
with its 2 genera and 5 species, contrasting strongly with our 
8 genera representing 11 species. A still greater difference is 
found in the Vitaceae, the European Vitis supporting but 2 
genera and 2 species, while our American vines afiford sustenance 
to 12 genera represented by 15 species. The European fauna 
of the Compositae is also much less, namely some G7 species 
representing 10 genera as compared with our 118 species assign- 
able to 15 genera. This large discrepancy is accounted for in 
great part by the enormous fauna of the Solidago and the num- 
erous species occurring upon aster ; plant genera which in Europe 
support only one genus and one species. 


J. P. Jensen. 

In several groups of animals, we find that the spermatozoa 
are held in packets or masses, and in some such as the Cephalo- 
pods, there is high specialization of the spermatophore, as the 
organ is called, in which the spermatozoa are contained. 
Among the insects, only representatives of a few groups form 
spermatophores, but when present they are beautiful and inter- 
esting structures and those formed by the gryllids or crickets 
are especially so. 

While engaged in research work on crickets in the Entomo- 
logical Laboratory of Cornell University, I noticed one day that 
a female Gryllus had a small pear-shaped organ attached 
between the ventral surface of the base of the ovipositor and 
the posterior end of the 8th abdominal sternite (Fig. 1). This 
structure did not seem to be part of her own body and as I did 
not at the time know anything about spermatophores, I was 
very much puzzled by it. After investigating the literature I 
found that I had happened to collect the specimen before the 
spermatophore had dropped ofif, and upon examining several 
males, I found a similar organ in situ in most of them, just out- 
side of what was considered the genital opening and covered 
by the ends of the posterior sclerites. 

Crickets, such as members of the genera Gryllus, Nemobius 
and Oecanthus are very difficult to classify and I had encoun- 
tered considerable difficulty in obtaining good specific charac- 
ters. More with a view of determining whether the spermato- 
phores might not assist me in classifying them, than expecting 
to make any morphological discoveries, I commenced to study 
them somewhat thoroughly. 

The literature was searched for accounts of copulation in 
these and related insects and five references were found. Ser- 
ville stated that in copulation the female Gryllus mounts the 
body of the male, as in the Oecanthids. Peytoureau said that 
in the Locustidae the transfer of spermatozoa takes place by 
imeans of a pear-shaped spermatophore that is transferred to the 


64 Annals Entomological Society of America [Vol. IV, 

female in copulation. Packard mentioned that in the two fami- 
lies of Gryllidae and Locustidae this was true and that especi- 
ally Gryllus had been noticed to have this habit. Gillette in 
1904 gave an excellent illustrated account of the structure and 
transfer of the spermatophores of the Western Cricket, Anabrus 
simplex, family Locustidae, and the most complete was found 
to be an account by Lespes in 1855, who not only noticed care- 
fully the complete process of copulation but b}^ dissection he 
determined how the spermatophores were developed and to 
some extent the function of the parts of the spermatophore. 

After relaxing some of my specimens of Gryllus, I dissected 
out the spermatophores in several males to gain a good knowl- 
edge of their structure. The spermatophore proper (Fig. 2, A) 
is attached to a handle-shaped part (Fig. 2, B) possessing five 
lateral hooks, three in front and two behind. The function of 
these hooks was not understood at the time but will appear 
later. A long whip-hke part (Fig. 2, C) is attached to the dorsal 
side of the handle. Many males were examined and a sperma- 
tophore was almost always found present. This is in accordance 
with Lespes' observations, who found that a new spermato- 
phore was completely formed in about one hour and that each 
female copulated several times during the egg-laying season. 

To determine the function of the parts, the female that had 
this organ still attached was after relaxation carefully dissected. 
The function of the hooks on the handle was readily found to be 
for attachment. The anterior part of the handle was found to 
be inside of the vaginal opening and the three anterior hooks 
held it firmly in place. The two at the posterior end also curve 
up and serve to hold it firmly in place by clasping to some extent 
the basal part of the ovipositor. After removing the bulb of 
the spermatophore, I attempted to remove the handle, but the 
anterior hooks held too firmly, part of it broke off but the whip- 
like structure remained attached and when pulled out, showed 
that it had extended a considerable distance up the passage 
and as will soon be shown this would indicate that the sperma- 
theca is quite far removed from the external opening. By 
mounting in glycerine and using high power the true relation of 
the whip-like part to the handle was made out, and also the 
nature and function of the former structure. It is attached 
somewhat nearer the spermatophore body than the middle of 

1911] Spermatophores of Crickets 65 

the handle and is continued as a dorsal thickening of the handle 
into the narrow cylindrical attachment between the handle and 
the bulb. It is in fact a duct, whose cavity can be traced from a 
point some distance inside the bulb (Fig. 2, D) to its outlet at 
the end of the whip. 

This was proven by embedding the tiny structure in paraffin 
and taking microtome sections of it (Fig. 3 and 4), from the far- 
ther end of the handle to almost the tip of the thread. The 
outside wall is rather gelatinous and soft, but a cylindrical, 
central core (Figs. 3 and 4, B) of very hard, apparently chiti- 
nous, material has the tiny duct in its center (Figs. 3 and 4, C) 
and in the sections this duct had not been flattened in the least. 
The very firm walls are no doubt for the purpose of preventing 
flattening or deformation, which might compress the duct 
and prevent the passage of the spermatozoa. . 

Last summer I again had opportunity to witness the courting 
and mating of Oecanthis fasciatus, and the process was very 
much the same as described by the writer in the Canadian 
Entomologist, Jan., '09. Then, however, I had missed the 
transfer of the spermatophore and after killing this female I 
removed the organ and mounted it in the usual way under a 
cover glass in Canada balsam. Watching it under low power 
of the microscope I succeeded by judicious pressure, in causing 
the spermatozoa to flow out of the end of the "thread. " This 
was final proof that this part of the organ is for conducting the 
spermatozoa to the spermatheca. Lespes in his account, some- 
how seems to have overlooked the fact that this thread-like 
structure is a duct, likely due to the minuteness of the duct 
itself, which when highly magnified, reminds one of a fine 
capillary tube. He calls it a horny thread, "file corne. " In 
fact he does not attempt to explain the structure at all, nor how 
the spermatozoa enter the vagina of the female from the sper- 
matophore after it has been placed in position. 

As before mentioned, the various species of crickets are very 
difficult to determine and th^ spermatophores may in the future 
be of considerable importance for definitely defining the species. 
For instance, Minnesota specimens of Gryllus pennsylvanicus 
Burm. vary considerably in general coloration and size from the 
Eastern specimens, but the spermatophores examined were all 
exactly alike. Lespes described and figured the respective 
spermatophores of the common European species, Gryllus 

66 Annals Entomological Society of America [Vol. IV, 

sylvestris, G. campestris and G. domesticus, and they differ 
very markedly from one another. Oecanthis fasciatus Fitch 
and O. quadripunctatus Beut. are the same species as gradations 
in the antennal markings show very nicely when one has consid- 
erable material. Whether the spermatophores further verify 
this, I have not as yet been able to definitely determine but it 
appears to me that the spermatophores of insects are worthy 
of considerable more attention than has been devoted to them 
in the past. 


All figures magnified, 3 and 4 highly. 
Fig. 1. Attachment of spermatophore to female Gryllus pennsylvanicus. A, bulb 

of spermatophore; B, Sth abdominal sternite; C, ovipositor. 
Fig. 2. Spermatophore of G. pennsylvanicus, magnified. A, bulb; B. handle; 

C, thread-like part (Lespes "file come"); D, duct; E, cross-section 

shown in Fig. 3; F, cross-section shown in Fig. 4. 
Figs. 3 and 4. Cross-sections of thread at E and F in Fig. 2. A, gelatinous 

outside wall; B, hard core; C, duct. 

Annals E. S. A. 

Vol. IV. Plate V. 

J. P. Jensen. 


Glenn W. Herrick. 

This is an European insect that is gradually becoming quite 
widely distributed in the northeastern United States and 
parts of Canada. It is also evidently causing considerable 
injury to larch trees wherever it is present. 

It was first noted in this country by Dr. Hagen, who, in 
1886, recorded it as seriously injuring the European larches on 
an avenue in Northampton, Mass. In 1905, Dr. Fletcher 
recorded its injuries to larches in Canada and in 1906 Miss 
Patch says that the case-bearers have been present in certain 
counties in Maine and " although minute they have been present 
in such enormous numbers that larch trees have often been, 
during the past three summers, eaten bare of green early in the 
spring." The insect has been present on the larches in the 
vicinity of Ithaca for several years, and undoubtedly does con- 
siderable injury every season. The small green leaves are 
devoured in early spring as fast as they push out, and on many 
trees the green tissues are eaten out and the leaves left pale 
and bleached in early spring. As soon as the buds begin to 
break in the spring, the dark brown, cigar-like cases that have 
been lying quietly attached to the branches all winter, become 
suddenly animated and commence crawling to the tender green 
leaves. In the spring of 1910 we found them active and feeding 
by the 16th of April. Each larva selects a leaf and soon eats 
a circular hole through the epidermis, thus gaining access to 
the tender tissues within. Then holding its case at right 
angles to the leaf and never releasing hold of its case it mines 
to the right and left of the opening as far as it can reach. The 
mined portion of the, leaf assumes a bleached appearance and 
the whole tree soon shows the effect of the injuries if the larvae 
are abundant. Observations would seem to indicate that the 
larvae molt just before leaving their winter quarters on the 
branches. This point, however, must await another season 
for definite determination. The cases of the larvae are enlarged 
after they have been feeding a few days by slitting the old case 
.and inserting a piece of leaf in the slit and fastening it in with 


1911] Life-History of the Larch Case-Bearer 69 

silk. One larva must attack a great number of the small 
young leaves, for in cases observed the larvae were not abundant 
enough to do the damage they did unless each case-bearer 
attacked and injured several leaves. As bearing on this 
point I selected a branch 6 inches long and found that it bore 
24 whorls of leaves, one whorl, at this particular stage, contain- 
ing 54 small leaves and other nascent ones in the center that 
could not be counted. If we take 54 leaves as the average, 
the branch bore 1296 leaves that were of a size to be attractive 
to the larvae. On this branch were 10 case-bearers. They 
had injured every leaf on the branch except those in the last 
whorl evidently having begun near the base of the branch and 
worked outward. These ten larvae had probably attacked 
and injured over a thousand leaves the majority of which, of 
course, were small. 

On April, 26th I found the first pupa in the breeding cages. 
When ready to pupate, the larvae attach their cases securely to 
the branches or to the leaves often in clusters of 4 or 5. A 
favorite place for attachment seems to be the center of a whorl 
of leaves. The period of pupation, in the breeding cages at 
least, proved to be from two to three weeks. We found moths 
emerging in the insectary May 11, 13, 15, 16 and on. 

The moths begin pairing in a few days after emergence 
and on May 31, their pinkish-red eggs were found deposited on 
leaves in the 'breedings jars. The eggs are shaped as though 
moulded in a tea-cup with many ridges radiating from the upper 
and smaller end, for they are glued to the leaves by their 

On June Gth, in the field, an abundance of moths were found 
and many of them were pairing. Some had probably emerged 
a few days earlier. On June 10th I found eggs on the leaves in 
abundance but there were still many pupae in cases showing 
that the moths emerge over a Jong period. The eggs are 
evidently placed indiscriminately on either side of the leaves. 

On June 2Sth and 29th the eggs were found hatching in the 
field. The egg-shells remain glued to the leaf and show no 
rupture of any kind for the emergence of the larva. Investi- 
gation shows that the larva bores through the base of the egg- 
shell and goes directly through the epidermis into the leaf 
beneath the egg. Here the larvae live mining in the tissues of 
the leaf but growing very slowly. The excrement of the tiny 

70 Annals Entomological Society of America [Vol. IV, 

larva is packed behind it in the mine. Here the larvae live 
until September. Owing to our absence from the University 
during the first part of September, we are unable to say at just 
what date the larvae first began to leave their mines and 
make their cases. On my return on the 15th of September 
many of them were found in their tiny cases feeding on the 
leaves. From this time on through September opportunit}- 
was given to observe them making their cases. In most 
instances, at least, they clean out their mines and pack the 
excrement in the outer end of it near the tip of the leaf. When 
the burrow is clean enough to suit them they cut off the tip 
of the leaf containing the excrement, which falls to the ground 
out of the way, and then they cut off enough of the leaf contain- 
ing the cliean part of the mine to make them a case of the desired 
length. The larvae now feed on the leaves of the larches until 
the latter part of October when they migrate to the branches 
and go into hibernation. 






By A. A. GiRAULT and James Zetf.k, 
Office of the State Entomologist of Illinois. 

In presenting for publication the results of a third successive 
year's observations on the biology of this insect made in the 
latitude of Urbana, Illinois and supplementing those made in 
Georgia in 1906 (Girault and Rosenfeld, 1907) and in Ohio in 
1907 (Girault, 1908), it becomes necessary to state that little 
or no progress has been made in regard to the continuity of 
observation and experiment, so that they should still be classed 
as desultory. The observations were made in the open or 
east insectary of this office at Urbana under as normal conditions 
as possible, but during odd hours and without previous fore- 
thought or planning and subject to much neglect at a critical 
time toward the last. 

They are presented, therefore, mainly to add to the sum of 
biological data on this insect, which in the end may lead to the 
discovery of important laws. At present, however, they form 
but a small beginning and cover but one or two biological fac- 
tors ; as they supplement to a large degree the observations made 
in Ohio (Girault, 1908), they are presented in the same general 

Those who gather data of this kind cannot help being 
impressed by our poverty in this respect and by the urgent 
necessity of accuracy in observation, to the minute as regards 
time and to the fraction of a degree as regards temperature, 
though it is true that such errors as occur should be chance 
errors, hence negligible. And most decidedly other factors 
should be taken into consideration, for in matters of this kind, 

* This may seem a trivial matter but consistency demands that the specific 
name of this insect be written as it was originally by Say; I see no necessity for 
change or reason therefor and certainly stability in nomenclature is not aided by 
making one. See articles 15 and 19, The International Code of Zoological Nom- 
enclature as Applied to Medicine (Stiles, 1905). If a change was necessary the 
form \-lineata would seem preferable to the other, being less radical. A. A. G. 


72 Annals Entomological Society of America [Vol. IV, 

we cannot foresee of what great importance the most trivial 
observations may become in the future and there is, doubtless, 
more than one cause for variability in periods of development. 


The following paper merely contains additional biological 
data along the same lines as those presented previously, obtained 
during the season of 190S, together with an account of the 
breeding of adults in confinement which resulted in reproduc- 
tion by the second generation of adults under adverse condi- 
tions. This reproduction by the second generation of adults 
apparently, was further hindered by actual starvation and was 
scanty, but the behavior of the beetles would lead to the belief 
that they were both willing and eager to reproduce. The fact 
is clearly shown that reproduction occurred with a pair of normal 
adults of the second generation, a result contrary to what we 
understand to be the meaning brought out by Tower (19()()), 
discussed previously (Girault, 1908). We do not, however, 
make any claims, but the evidence is sufificient to establish the 
fact that exceptionally the adults of the second generation in 
normal beetles do develop the germ-cells before a period of 


1. Length of Stadium* 

The duration of embryonic development was determined 
for about nine hundred cases during the breeding season and 
the results are tabulated in Table I. The separate lots were 
confined as previously, in darkness. In every case recorded 
the time is actual, unless noted to the contrary. By comparing 
these records with those given by Girault (1908, Table I, 
p. 156), differences are noticeable in regard to the duration of 
the stage at the same approximate dates for the two latitudes; 
witness Lot I of the two tables. We should expect to find here 
a corresponding difference in the temperatures. 

* This term is used in preference to instar whieli was originally proposed to 
designate the insect itself at any stage or perioil of development, as the egg instar, 
third larval instar and so on, just as we say the larva, caterpillar, pupa or imago. 


Notes on the Colorado Potato Beetle 


Duration of the Egg Stadium, Urbana, III., Season 1908. 








Effective Temp 
Daily Aver- 
ages, Degrees 









Pair No. 3 Hib. adults . . 

;; 1 Gen. I : ' 

.■) Hib. adults . 

1 Gen. 1 

" 3 Hib. adults . 
;] 1 Gen. 1 

3 Hib. adults '. '. 

1 Gen. 1 

3 Hib. adults . . 

I '^ I ■■ 

3 " '.'. 































11:45 a.m. 
1:30 p. m. 

7:00 p. m. 
10:45 a. m. 

3:00 p. m. 

1:00 p. m. 

1:15 p. m. 

3:00 p. m. 

4:30 p. m. 

1:00 p.m. 

1:45 p. m. 

4:00 p. m. 

2:30 p. m. 

2:30 p. m. 

3:15 p. m. 

3:.30 p. m. 
12:30 p. m. 
11:30 a. m. 
10:00 a. m. 
10:30 a. m. 
10:30 a. m. 




9:45 a. m. 
11:30 a.m. 

7:00 p. m. 

6:30 p. m. 
12:00 m. 

5:45 p. m. 

7:00 p. m. 

5:00 p. m. 

1:00 p. m. 

9:00 p. m. 
11:30. a.m. 

9:00 p. m. 

7:00 p. m. 

6-00 a. m. 

8:00 a. m. 

6:00 a. m. 

5:00 p. m. 
12:00 m. 

9:00 a. m. 

2:00 p. m. 

3:00 p. m. 

















24 4° 

















But first attention siiould be drawn to the fact that there 
exists variation in the duration of embryonic development for 
batches of eggs deposited at the same time, hence subject to the 
same environmental factors including temperatures. Thus in 
lots 13 and 14 (Table I), from different parents, deposited at 
the same time on June 14 hatched at different times on June 
19, lot 14 hatching 13 hours earlier than lot 13. And in 
lots 8 and 9 ; although there is a difference of an hour and a half 
between the times of deposition, the times of hatching diverge 
still more being separated by four hours and the lot deposited 
last hatched first. These lots were from the same parent. But 
contrary to this, in lots 10 and 11 deposited by different parents 
within 45 minutes of each other, the lot deposited first hatched 
first, the times of hatching being 141 hours apart. However, 
lots 20 and 21 deposited by different parents at the same time 
hatched within an hour of each other. The data are insufficient 
but parentage apparently does not account for the variation 
between batches of eggs deposited simultaneously and we must 
state tentatively that it is inherent and hence subject to the 
laws of chance or else there are factors involved which have 
escaped detection. We think this variation is inherent and 

74 Annals Entomological Society of America [Vol. IV, 

hence limited or continuous and with sufficient data could be 
plotted in the same way as other continuous variations. It is 
of the same nature, apparently, as individual variations in the 
duration of postembryonic stadia, a matter of common obser- 
vation and which are not controlled b}^ temperature within 
certain time limits, nor by food. 

As found previously, the daily average effective temperature 
increases as the period of embryonic development decreases and 
conversely. But for equal periods of development as shown in 
foregoing, equal amounts of temperature were not necessary, 
as witness lots 1 and 2, 7 and 12 and lots 13 and 21; also lots 
9, 11 and 19. For a degree of temperature (effective) there 
appears to be a variable amount of growth or development, 
which as yet remains unpredictable; it is a specific, or maybe 
generic, characteristic. 

2. Number of Eggs Deposited. 

The data obtained on this point but serve to confirm what 
is stated by Girault (1908, p. 157 ff.) in a previous paper and 
also to increase the maximum number observed to be deposited 
by several hundred. The data were derived mainly by keeping 
in confinement three pairs of hibernated beetles captured early 
in the season while mating in a potato field and one or two pairs 
of the succeeding generations. The total number of eggs 
deposited, the rate of deposition and other related points for 
the pairs of the several generations are brought out in Table II 
presented herewith. The records fall short of what actually 
would have been the totals for the generations, as toward the 
second week in August the adult beetles were much neglected 
and finally died of starvation. The effect of this lack of nour- 
ishment on the second generation (or parents of the third gener- 
ation) was especially noticeable, for although mating occurred 
freely throughout the different lots, oviposition occurred but 
once and most of the beetles disappeared into the soil for hiber- 
nation nearly as soon as their food was discontinued. The 
results indicate, however, that the first generation of adults are 
capable of as large an amount of reproduction as are the hibernated 
beetles and that the second generation of adults (or parents of 
the third generation) were willing or able to reproduce. 

The three pairs of the hibernated beetles were obtained 
from a potato field in Urbana captured while mating at 11 

TABLE II. Numb; 




Pair No. 1. 





7:00 p, m. 




7:00 p. m. 




2:00 p, m. 




— : — a. m. 




— : — a, m. 




7:00 p, m. 




12:00 M. 




— :— a, m. 




11:00 a. m. 




10:45 a. m. 




3:00 p, m. 




3:30 p, m. 




2:00 p. m. 




4-6 p, m. 




3:00 n, m. 




1:30 p. m. 




9:00 p, m. 




2:30 p. ni. 




1:00 p, m. 




4:00 p. Ill, 




3:00 p, m. 




10:00 a. m. 




1:30 p, m. 




2:45 p. m. 




2:,'iO p. m. 





























of batches 


per batch 

Daily av 



TABLE II. Number of Eg(;s Deposited in Confinement by Pairs of Different Generations, 1908. 


Hibernated Adults. 

Generation I. Parents 2d Gen. Generation II. Parents 3d Gen. 


1st Mating observed: 


First mating observed 
June 23, 9 a. m. 

First mating observed 



23, 11 a. 




May 27, 7:30 p. m. 

Pair No. 3. 


1 , 




Aug. 11, 3 p. m. 

Pair No. 1. 


Pair No. 2. 

Pair No. 1 


Pair No. 2 






Date 1 



May 23, 7:00 p. m. 


May 23-27 


May 28, 1:30 p. m. 


Julv 8.—:— a.m. 

52 1 

Aug. 11, 10.00 a.m. 



" 30. 7:00 p. m. 


" 30.11:00 a.m. 


" 29,11:45 a.m. 

58 , 

S, 3:00 p. m. 

58 1 

" 12, — : — p. m. 



June 1, 2:00 p. m. 


" 30, 5:00 p. m. 


" 29, 3:00 p. m. 


" 8, 4:30 p. m. 

43 1 


" 3, — : — a. m. 


June 1, 1:00 p.m. 


" 30, 1:30 p.m. 


9. 2:00 p. m. 



" 4, — : — a. m. 


2, 2:00 p. m. 


Junt' 1, 1:30 p. m. 


" 10. 1:00 p.m. 



4, 7:00 p. m. 


4. 12:00 M. 


2, 2:30 p. m. 


" 10, — :— p. m. 


5, 12:00 M. 


5,10:30 a.m. 


4, 2:00 p. m. 


" 11, 10:00 a.m. 




" 6, — :— a. m. 


5. 10:00 a. m. 


" 11, 4:00 p. m. 



" 7. 11:00 a. m. 


6,—:— a.m. 


6, 11:00 a. m. 

36 1 

" 12, — :— a. m. 



" 8, 10:45 a. m. 


7,11:30 a.m. 


" ' 6, — :— p. m. 


" 13, 1:00 p. m. 



9. 3:01) p.m. 
'• 11, 3:30 p.m. 
" 13 2:00 p. m. 
" 14, 4-6 p. m. 
" 17 3-00 T) m. 


8, 12 M. 


7.11:30 a.m. 


" 14, 2:30 p. m. 




9, 3:.30 p. m. 


7, 7:30 p. m. 


" 15. 12:30 p. m. 




" 10, 3:00 p. m. 


8, 12:.30 p. m. 

39 ] 

" 16. 8:00 p. m. 80 1 


" 12. 1:00 p.m. 


9.2:30-4:30 p.m 


" 17. 12:00 M. 




13, 3:01) p. m. 


" 10, 4:00 a. m. 


18. 3:15 p. m. 



" 22, 1:30 p.m. 
'■ 23, 9:00 p. m. 
" 24, 2:30 p. m. 
" 26, 1:00 p,m. 
" 26, 4:00 p. m. 
" 28. 3:00 p. m. 
" 30. 10:00 £. m. 


" 14. 10:00 a. m. 


11, 3:45 p. m. 


19, 12:30 p. m. 



20, 3:00 p. m. 


" 12, 1:00 p. m. 


" 21. 11-30 a.m. 



" 21, 1:20 p.m. 


" 13, 2:45 p. m. , 


" 21-22, a. m. 



" 22. 1:30 p.m. 
" 23. 9:10 a.m. 
" 24. 1:30 p.m. 
" 28. 2:20 p. m. 


" 14, 4-6 p. m. 
" 16. 1:30 p.m. 
" 17, 
" 21, 1:30 p. m. 

31 i 

1 i 

41 1 
28 1 

" 24, 2:00 p. m. 
" 25. I0:.3I) a. m. 
" 26, 12:00 M. 
" 27. 10:00 a. m. 




July 1, 1:30 p.m. 

" 3. 2:45 p. m. 

4, 2:50 p. m. 


" 29, 3:00 p. m. 
July 1. 10:00 a. m. 

2, — :— p. m. 

3. 2:40 p. m. 


" 22, 1:30 p. m. 

" 23, 1:10 p. m. 

" 26, 5:30 p. m. 

July 4, 3:30 p. m. 

6, 1:20 p.m. 


6, 1:30 p.m. 



7. 3:00 p. m. 


8, 2:00 p. m. 


9. 11:45 a. m. 


9. 1:30 o.m. 


" 10. 1:00 p.m. 


10, 1:45 p. m. 


12. 9:30 a. m. 


« 11, 2:00 p.m. 


" 12. 1:00 p. m. 


" 12, 9:00 a. m. 


« 14 2:30 p. m. 


" 14, 2:30 p. m. 


" 15. 12:00 M. 


" 14, 5:00 p. m. 


" 17. 11:45 a. 'm. 


" 15, 3:00 p. m. 


' " IS. 10:30 a. m. 


18, 10.00 a. m. 



19. 1:30 p.m. 


" 19, 3:30 p. m. 


" 21. 1:00 p.m. 


" 23,11:30 a.m. 


" 25, 10:00 a. m. 


" 25, — :— p. m. 


" 25. 10::!0a. m. 


" 26, 4:00 p. m. 


Aug. 2, 
" 3 


" 29, — :— p. m. 
« 30, 10:00 a. m. 




4. 10:00 a. m. 


Aug. 2, 


5. 10:30 a. m. 


3, — :— p. m. 



" 6. — :— p. m. 

; " 7,11:00 a.m. 

" 11, 10:00 a. m. 




4, — :— p. m. 

5, 10:30 a. m. 
" 6.—:— p.m. 




" 16. — :— p. m. 


8, — :— p. m. 




" 10. — :— p. m. 
12, — :— p. in- 

1 1 




1 . — 



Total KI-1" 


No. of batches 1 -'2 

Av. per batch ; 47^ 

DailvAv 1 52 

Tot,, 13 

Total ^ . 

1 Total 

Average, 1280.6 


No. of batches 

No. of batches 

Av. per batch 

36 08 

No. of batches 48 

•■^v. per batch i 28.37 

nii)v TV . . . . ! 16 40 

1 No, of batches 

1 Av. per batch 

1 Daily av 


Av. per batch , ■• 

1 Daily av ' « 

Daily av 


1911] Notes on the Colorado Potato Beetle 75 

A. M., May 2.3 (pairs No. 1 and 2) and at 7:30 P. M., May 
27, 1908 (pair No. 3) and confined with food immediately 
after capture. The single pair of the first generation resulted 
from a mass of 60 eggs deposited by hibernated beetles and 
taken from the field on May 23, 1908 and the single repro- 
ducing pair of the second generation are direct descendants 
■of the pair of the first generation. 

In the case of an extra cage containing a large number of 
adults collected in the field during the latter part of July, a 
female was observed to deposit a mass of 103 eggs, the largest 
single mass of eggs yet recorded. In another case, the rate of 
•oviposition was timed; a female deposited in succession in a 
single mass in the usual manner 64 eggs in a period of time 
occupying 3200 seconds or 53 1-3 minutes. The rate of deposi- 
tion was regular, each single, deposit requiring 50 seconds — 40 
seconds to pass the egg and to fasten it and about 10 seconds to 
obtain position for the next deposit. 

Attention is called to the rapid depo.sition of the single 
pair of the first generation, having a daily rate of deposition 
of 52 eggs and on a single day (July 8) depositing as many 
as 153 eggs in three separate batches, averaging 51 eggs each. 


1. Duration of Larval Stadia. 

We were able to make more observations concerning this 
phase of the beetle's life during 1908 than at previous times. 
The records for the first fifteen lots in the annexed table (Table 
III) comprise single larvae of the same age and parentage, that 
is, they are all from the same batch of eggs, hatching at the 
same average time but confined separately each individual 
ecdysis being recorded. 

Lot No. 16, comprising 45 larvae, was from the same mass of 
60 eggs as the larvae of lots No. 1-15, but upon hatching were 
confined together on their food. With them, the first ecdysis 
became general at 4 p. m.. May 29; the second ecdysis began 
at 7 p. M., May 31, but was not general until 2:30 p. m., 
June 1, and was completed at 6 p. m., June 1, occupying 
a period of 23 hours. On June 3, the larvae were large, 
plump and healthy, eating voraciously, but only 30 in number, 
15 having died. The third ecdysis began at 5 P. M., June 4, 












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■ ^ • .CO 



■ - • -POCQN • "mm ■ - -CCMM 


■ •■^ ■ -eo 

Entered soil 
Stadium IV 




; ; ; eee ; ee ; e 

■ ■- d.rtrt- doi- ctii 

■■'„...:<".. :q5 t,' . 
: : :g : 

... 3 - 3 33 





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a 35 




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2' :s s; c:Sa>-^ c<.:ss;s;,o 


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dD.dddD.d.d.rtajc.Q.D.c3 dci 
iOOOco O eoOO e^oeo O O O cc O 





B .6E 

rtS dd 


^ — M — I- — <D —■ — O: M M •-■ t~ M SO 

—* —'■ — T — " — T _h' — ' — " — " ^" -J —*.-.' ^ -^ erf 
c* CO M CO CO CO eo ec CO eo — 

(U >. CJ Oj >, 

1 i 



— MM -MCM -MCOMMu:)— W t- 


— lO — MCO 



MCg^ — M — MM — M — MMMMMCO 







EE .E 

cjddrtdndreaidajddD. eft 

gsggggggglSggs gg 

— oir^t^ — » — — ciJMO — COM "^M 



o. a.<^ o. 1 











g g.gSfe 

. . . .< 

„ CO'J'CD*^ 

= = . = <« 

,,,.=== s^" 



E-i -ON 


S 3§S 

"^ "" .<.!.- 


3N -ioi 


Mco'a*ioor-ooeso^M«'*»o -(ot^oo 

O T3 dji— 
t-5 MMM 


Notes on the Colorado Potato Beetle 


and was general at 8 A. M., June 5, concluding at 2 p. M., 
June 5, occupying a period, of 21 hours. But 18 larvae suc- 
■cessfully survived the ecdysis. Entering the soil for pupation 
began on June 7, at 4:30 a. m. and all larvae had entered 
by June 8, 7 p. m. Table III summarizes. 

2. Number of Ecdyses. 

There can be no doubt but that the normal number of 
larval ecdyses, excluding pupation, is three and as additional 
evidence we have observed this number in two hundred and 
the fifty cases during the season without a single exception for 
whole number. The question may be considered as settled. 

3. Duration of the Larval Stage. 

There being no data concerning this point other than what 
are already included in Table III, it is tmnecessary to repeat 
them here, but reference should be made to the column of 
sums of that table. 


1. Duration of Pupal Stage. 

Table IV summarizes sufficiently well all of our data for 
1908 concerning this phase of the beetle's life cycle. 

Duration of Pupal Stage, Actual Time in vSoil, Season 1908. 


' Length Time 
in Soil. 



Entered Soil. 

Adults Emerged 

Sum of Effective 


Temp. Degrees 





June 7. 4:30 a. m. 


21, 1:30 p.m. 





8, 7:00 p.m. 


22. 6:00 a.m. 





" 7. 4:30 a. m. 


19, 4:00 p. m. 





7. 4:30 a.m. 


21. 1:30 p.m. 





8. 7:00 p. m. 


22, 1:00 p.m. 





7, 4:30 a.m. 


21. 1:30 p.m. 






" 7, noon 


22. noon 




July 19, 7:00 a. m. 


30, 6:00 p. m. 





24. 8:00 a.m. 


4, 7:00 p.m. 






25. 9:00 p. m. 

6. 7:00 a. m. 






24.11:00 p.m. 


4. 8:30 a.m. 






25,11:00 p.m. 


5. 12:15 p. m. 






27. 6:00 a.m. 

0, 6.00 p. m. 




• These numbers correspond with the lots in Table III. 
t Average of Lots No. 1 to 12, 13 days, 193^ hours. 


Annals Entomological Society of America [Vol. IV, 


1. Length of Life in Confinement. 

a. In Pairs Normally Reproducing. 
The data obtained on this point are scanty and much 
vitiated by the fact that the lots were neglected too soon to 
obtain normal results, but they supplement to some extent the 
data obtained in 1907 tending to support the theory that the 
average duration of life of normally reproducing adults is two 
months or more. The average here is 1.8+ months, the data 
however being insufficient. 

Length of Adult Life in Confinement, Normally Reproducing. 

Lot No. 



Date Confined. 1908 

Date of Death. 1908. 

Length of Life, 







I. Hibernated 




II. Gen. I 


III. Gen. II 








Potato field, mating. 

Hibernated adults 

Pair No. 1, Gen I. 

11 a. m., May 23 
11a.m., May 23 
7:30 p.m., May 27 

June 23 

July 30— Aug. 8 

June 4" 
Aug. 16t 
July 26 

Aug. let 


July 7 
Aug. I6t 
Aug. I6t 

Aug. 16t 







1 5 




* Escaped. 

t Liberated. 

; Starved and entered hibernation. 

2. Length of the Period of Oviposition. 

As with the previous section, the results here are abnormally 
short in point of duration for the reasons given. They are 
merely tabulated therefore, without further comment. 

Length of the Period of Oviposition. 
Generations, 1908. 


Generation No. 

First Mated. 

First Eggs 

Last Eggs 

Length of Period of 
Oviposition, Days. 

Hibernated — 

Pair No. 1 

11 a. m.. May 23 
11 a. m.. May 23 
7:30 p.m.. May 27 

7 p. m.. May 23 

May 25' 

1:30 p.m.. May 28 

2:50 p.m., July 4 
August 16, p. m. 
August 12, p. m. 

42 + 

Pair No 2 




I— Pair No. 1 

9 a.m.. June 23 

July 8. a. m. 

10 a. m., July 27 


II.— Pair No. Ic 

Aug. 11, 3 p. m. 

Aug. 11. 10 a. m. 

Aug. 12. p. m. 


* Average time of a period of 4 days. 


Notes on the Colorado Potato Beetle 


3. Mating. 

The observations on this habit are also Hmited, but those 
matings actually observed are summarized in Table VII. 
In a single case, the time actually involved from beginning to 
end of the act was obtained, being three and one-half hours 
(10:30 A. M. to 2 P. M., June 18, Pair No. 3, hibernated adults.) 


Frequency of Mating in Reproducing Pairs. 

Different Generations, 1908. 




First Mating 

Subsequent Matings 

Last Matings 


wa a 










11 a. m.. May 23 

/May28. 30. 31 \ 

IJune 1 ; 

June 4* 



42 + 



11 a. m., May 23 

May 27. 30 

June2. 4. 21 

ijuly 1, 23, 26. 28. ... 

lAug. 3. 4, 5. 5 

[ 7.7 

August 11 






7:30 p. m.. May 27 

[May 30 

July IS 



June3. 4. 10. 13, 18. 


(July 1, 9, 15, 17. 17 . 




9 a. m.. June 23 

July 8 







Aug. 3. 3 p. m. 
Aug. 9, 4 p. m. 
Aug. 9. 10 a. m.° 
Aug. 9, 10 a. m° 1 
Aug. 9. 6:30 a. m.°J 

Aug. 4. .5. 6 

Aug. 11, 12 

Aug. 7, 9 a. m.t 
Aug. 13. 3 p. m. 







Aug. 11° ! 
Aug. 11 ) 

Aug. 13 

Aug. 14 



* Male escaped. t Male died July 26. t Male entered soil for hibernation. 
° Only observed mating; hibernation followed within 10 days. 

Mating was observed during the following hours of the day: 
Practically at any hour between 7 A. M. and 11 P. M., more 
commonly at 9, 10 and 11 A. m. and 1, 2, 3, 4 and 6 P. M. or 
at fractions of those hours. The function was observed most 
commonly at 9 and 10 A. M., over 31 per cent of the 58 times 
the act was observed being either at or between those two 
hours. Fifty per cent of the observed matings occurred in the 
morning and fifty per cent, in the afternoon or evening. Ob- 
servations were continued throughout most of the night, up 
at least until midnight, commencing again at six o'clock in 
the morning. 

80 Annals Entomological Society of America [Vol. IV, 

4. Potency of Fertilization. 

As concerns this point, it was noticed in the case of the hiber- 
nated pairs, and with these pairs only was opportunity presented 
to gather any data bearing on the questioh, that the female of 
Pair No. 1 continued to deposit fertile ova for one month' after 
the absence of the male (June 4, 7 P. M., to July 4, 2:50 
p. M.) ; and that the female of Pair No. 3 deposited fertile ova 
for seventeen days after the death of her mate. No other 
data were obtained. 

5. Number and History of Generations Reared in the Laboratory. 
Our data here are also meagre, but they certainly do tend to 

uphold the opinion that the adults of the second generation 
(or parents- of the third generation) are at least able, if not 
willing, to reproduce and hence the observations of last year 
(Girault 1908) are upheld and Tower's (190G) dictum that 
"The second generation does not develop the germ-cells nor 
show any reproductive activity until after it has passed through 
a period of hibernation or aestivation" becomes in our minds 
less and less authoritative. These beetles of the second gen- 
eration with us certainly showed reproductive activity, if 
repeated matings can be called such, and one pair, even under 
very adverse conditions — starvation — deposited fertile eggs, 
which surely must be conceded to be reproduction which cannot 
of course take place without development of the germ-cells. 
The beetles with us this year plainly showed symptoms of what 
we would call eagerness and ability to reproduce. These beetles 
were those of the second generation, as will be shown in the fol- 
lowing brief historical sketch, and were normal in everj^ way, 
that is to say, did not represent any special race of the species. 
On May 23, 1908, or at the earliest possible date, 60 eggs 
of the species were collected from a potato plant in a small plot 
of potatoes at Urbana, Illinois and brought to the laboratory 
to comprise the first generation or descendants of the hiber- 
nated adults. The larvae came to maturity early in June and 
pupated and eleven adults emerged between June 21 and 
June 23. They were confined together with food. On the 
latter date- a pair were- found mating and were at once isolated 
as the parents of the second generation.* From this pair of 
adults of the first generation, there were taken for the special 

*The others were accidentally poisoned with arsenate of lead. 

1911] Notes on the Colorado Potato Beetle 81 

purpose of rearing a sufficient quantity of the second generation, 
six lots or batches of eggs numbered from a to/. In all 49 adults 
were obtained from the six batches. For clearness, the batches 
are treated in detail: (1) Batch a, consisting of about 20 
(number unknown) eggs hatched at 6:30 A. M., July 7, the 
resulting larvae entering the soil for pupation at the average 
time of 7 A. m., July 10 and on July 30 and 31, 4 adults 
were obtained. These were at once fed and at 3 p. m., August 
3, a pair were observed mating and were isolated. This pair 
continued to mate until A. m., August 7, the c7 entering the 
soil shortly afterward; with them mating was observed five 
times, but no oviposition occurred. In the meantime, the 
two remaining beetles had hibernated (August S), the mated 
female following a week later. Hibernation induced by star- 
vation due to lack of time in which to feed the beetles. (2) 
Batch b, consisting of 60 eggs came to larval maturity at S A. M., 
July 24, and on August 4 and 5, two adults were obtained 
comprising the whole survival. These were males and hiber- 
nated on August 22. (3) Batch c, 58 eggs, came to larval 
maturity at 9 p. m., July 25, and gave from August 5 to 
7, 4 adults which were placed on food as they emerged. A 
male died on August S and a pair were mating at 4 p. m., 
August 9; this pair was then isolated. On August 11 at 
10 A. M., 3 eggs were deposited which proved to be fertile; 
mating was again observed at 3 p. m. the same day and at the 
same hour on August 12; later the same day (12) 9 eggs 
were deposited on a leaf, which also proved to be fertile; another 
mating was observed at 9 A. M., August 13, but thereafter no 
other matings were observed and further reproduction did not 
occur. The remaining adult died on August 22, but the 
mated pair remained aUve without food until August 25, 
when the cage was broken up. Oviposition and mating in 
spite of insufficient food. (4) Batch d, 43 eggs, came to larval 
maturity at 10 i'. M., July 24, and on August 4 from 7 to 
10 A. M., 11 adults were obtained, the total survival. On 
August 9, 3 pairs observed mating were isolated but other 
matings did not occur with them nor oviposition, caused as we 
have reason to believe, by the neglect to supply food. Thus, 
on August 13 the third pair had entered the soil for hibernation 
and two days later the second pair had done likewise; 
the first pair remained on top of the soil until August 25, 

82 Annals Entomological Society of America [Vol. IV, 

when they were killed and removed. Of the remaining five 
adults, two had died by August 11 and the three others hiber- 
nated on August 22. (5) Batch e, 56 eggs, came to larval 
maturity at the average time of 11 p. M., July 25, and gave 

24 adults from August 4 to August 6, which were confined 
together with food. But a single pair was isolated, observed 
mating on August 1 1 , though previously, mating had occurred 
promiscuously. This pair was neglected after isolation and 
no further reproductive activity occurred; on August 13 at 
9 A. M., the male entered the soil for hibernation and on August 

25 the pair were removed still alive. Of the remaining 22 
beetles, 4 hibernated on August 11 at 9 a. m. and by August 
22, all had disappeared beneath the soil, two having died 
there. No reproduction, but during the period of feeding, after 
several days, mating was frequent and promiscuous and there 
is good reason for believing that reproduction was prevented 
by actual starvation at a critical period. (6) Batch/, 32 eggs, 
arrived at larval maturity at the average time of 6 A. M., July 
27, and gave 4 adults August 7 and 8; on August 11, 
a mating pair of this lot were isolated and the remaining two 
also paired. The first pair mated again on August 13 and 
August 14 but no oviposition followed and they were removed 
on August 25, after days of starvation. The second pair 
had hibernated by August 22, without depositing eggs and 
with no further observed matings. 

In general it may be stated that the adults of the second 
generation just after emergence fed voraciously for several 
days and then began to mate as though eager to reproduce and 
one pair acutally deposited fertile eggs, insuring at least a por- 
tion of a third generation. It was at this time in their lives, 
just following the period of heavy feeding and the beginning 
of mating that stress of other work caused the food to be neg- 
lected and after August 8, the beetles were starving and were 
forced to hibernate. Incidentally, it was also true that their 
food-plant in nature was also very scarce at this time, so even 
if at large, it is not unreasonable to suppose that these beetles 
of the second generation would have been forced into hiberna- 
tion before reproduction could begin, though willing and able 
to reproduce. What little evidence we have gathered this year 
forces us to conclude that the second generation of adults 
exceptionally are both willing and able to reproduce, merely 


Notes oil the Colorado Potato Beetle 


supplementing what was previously indicated to be true in 
1907. The evidence of course is gross in nature, for we did 
not actually examine the mated females in any case for sperma- 
tozoa, so that in the majority of cases, actual mating is open to 
question. It is needless to say that this should have been done. 
But in at least one case we are sure that both mating and repro- 
duction occurred as fertile ova were deposited. 

In regard to the seasonal history in 1908, the second genera- 
tion was obtained nearly a month earlier than that obtained 
in 1907, so that there was ample time for a third generation. 
The following table summarizes the generations reared in con- 

Generations Reared in the Laboratory, Urbana, 




Eggs Deposited. 

Adults Out. 

Length of Cycle. 





Temp Sums . 
Degrees F. 


May 21* 

June 22 



II. Lot a 

July 2t 

July 6, 1:1.7 p. m. 
July 8. 3 p. m. 
July 8, 4:30 p. ra. 
July 10. 1 p. ra. 
July 11, 4 p. m. 

July 30, 11 p. m. 
August 4, 7 p. m. 
August 6, 7 a. m. 
August 4, 8:30 a. m. 
August 5, noon. 
August 7, 6 p. m. 






August U and 12. 

Not reared to maturity 

■ Approximated; hatched 11 a. m., May 27. 

t Approximated; hatched 6:30 a. ra.. July 7. 


1905. Stiles, Charles Wardell, Bull. No. 24, Hygienic Laboratory, Public Health 

and Marine-Hospital Service of the United States, Treasury Department, 
Washington, D. C. 

1906. Tower, William Lawrence. An investigation of evolution in chrysomelid 

beetles of the genus Leptinotarsa. Publication No. 48, Carnegie Institu- 
tion of Washington, Washington, D. C. 

1907. Girault, Alexandre Arsene and Arthur H. Rosenfeld. Biological notes on 

the Colorado potato beetle, Leptinotarsa decemlineata (Say), with 
technical description of its stages. Psyche, Cambridge, Mass., XIV, pp. 

1908. Girault, Alexandre Arsene. Further biological notes on the Colorado 

potato beetle, Leptinotarsa decemlineata (Say), including observations 
on the number of generations and length of the period of oviposition. 
Annals Ent. Society of America, Cokmibus, Ohio, I, pp. 155-17S. 


The Fifth Annual Meeting of the Entomological Society 
of America was called to order at 10:45 A. M., December 27, 
1910, in the School of Mines Building, University of Minnesota, 
Minneapolis, by the President, Dr. John B. Smith. In the 
absence of the Secretary, Professor J. G. Sanders was elected 
Secretary pro tern. Announcements. 

Professor F. L. Washburn moved that the chair appoint a 
committee of three to confer with a similar committee from the 
Association of Economic Entomologists concerning the organiz- 
ation of an Entomological Employment Bureau or Clearing 
House. It was agreed that the organization of such a body 
would facilitate the securing of available men for entomological 
work. Several expressed favorable opinions concerning this 

The following papers were presented: 

Notes on the Tingid Leptobyrsa explanata Heid., by E. L. 
Dickerson; read by the Secretary. 

Notes on Sanninoidea exitiosa by Dr. J. B. Smith. Discus- 
sion by Mr. R. L. Webster, asking if any tables of head widths 
of various larval instars of this species had been published. He 
reported that such measurements constituted a very good 
method of identification. 

"The Structure of Spermatophores in Crickets," by Mr. 
J. P. Jensen. Read by the author. (PubHshed in March 

Dr. Smith asked if studies had been made of the copulatory 
organs in various species. Mr. Jensen replied that comparative 
drawings of a large number of individuals of the same species 
had been made, likewise of different species. He also reported 
that Nemobius fasciatus var. vittatus had been found in large 
numbers digging in loose soils, securing and destroying eggs of 
Melanoplus bilineatus. He considered this insect as undoubted- 
ly a considerable factor in the control of Melanoplus. Dr. 
Smith questioned: "Is not such the general habit of some 
Orthoptera?" Was answered by Professor Bruner, "Many 
Orthoptera are largely carnivorous." 

Professor Oestlund invited members to visit and inspect 
his collection of Aphididae. 


1911] Minutes of the Miiineapol is Meeting 85 

The Society then adjourned until 1 :30 P. m. 

The President appointed the following committees when the 
Society reconvened: 

Committee on Employment Bureau to confer with similar 
Committee from the Association of Economic Entomologists: 
Messrs. F. L. Washburn, Herbert Osborn, and Henry Skinner. 

Nominating Committee: Professors E. D. Sanderson, H. E. 
Summers and R. L. Webster. 

Auditing Committee: Professors Lawrence Bruner and J. G. 

The following papers were read: 

"The Biological Survey of the Insect Life of Kansas" by 
Professor S. J. Hunter. 

"An Experimental Study of the Death-Feigning Habit of 
Belostoma {Zaifha) flumineum and Nepa apiculata Uhler," 
by H. C. and H. H. Severin. Discussed by E. C. Cotton with 
the remark that the weevil Apion segnipes which worked in 
border pea-pods in Tennsesee was unable to free itself from 
the pod but is released automatically by the sudden opening 
of the pod. The Apion when disturbed under such conditions 
does not feign death, but if handled later it feigns death. 

"Announcement of Further Results Secured in the Study 
of Tachinidae and Allies," by C. H. T. Townsend, Piura, Peru. 
This paper was read in part by the Secretary. (To be pub- 
lished in June Annals.) 

The ' ' Report of the Committee on Nomenclature ' ' was written 
l)y Professor T. D. A. Cockerell with H. T. Fernald and E. P. 
Felt and was read by the Secretary. After some discussion, 
Prof. H. E. Summers moved to receive the report, order it 
printed and consider it at a later date. Carried. 

The Society then adjourned until Wednesday at 9:00 a. m. 

At 9:00 A. M., December 28, the Society was again called to 
order by the President, Dr. Smith, and the following reports 
presented : 

The Report of the Editor of the Annals, Professor Herbert 
Osborn, was presented and on motion of Professor Lawrence 
Bruner, was accepted. 

The Report of the Auditing Committee on the accounts of 
the Editor was presented by Professor Lawrence Bruner and 
accepted. He also reported on the accounts of the Treasurer 
for the Committee and they were accepted subject to correction. 

86 Annals Entomological Society of America [Vol. IV, 

The Report of the Secretary of the Executive Committee 
was presented and accepted and is given in full later. 

The following paper was read: 

"Some Suggested Rules to Govern Entomological Publi- 
cations," by T. D. A. Cockerell, read by the Secretary. Sev- 
eral suggestions were made by Dr. Wolcott concerning entomo- 
logical publications, as follows: 

That it is the privilege of contributors to demand proof of 
their papers, but it is also obligatory that corrected proof be 
returned as soon as possible. Likewise, it was remarked that 
contributors could not expect manuscripts to appear in print 
on extremely short notice, as is frequently the case, but should 
expect their papers to take their turn. 

The Nominating Committee reported as follows for ofificers 
for 1911: 

President — Professor Herbert Osborn. 

First Vice President — Professor Lawrence Bruner. 

Second Vice President — Professor A. D. MacGillivr.w. 

Secretary-Treasurer — Professor A. D. MacGillivray. 

Additional Members of the Executive Committee: 
Professor J. H. Comstock Dr. W. M. Wheeler 

Dr. J. B. Smith, Dr. H. Skinner, 

Professor C. J. S. Bethune, Dr. A. D. Hopkins. 

It was moved by Professor M. H. Swenk that the Secretary 
be instructed to cast a unanimous ballot for the officers nomi- 

Professor E. D. Sanderson moved that a vote of thanks from 
the Society be extended to Professor Herbert Osborn, Manag- 
ing Editor, for his faithfulness and especial care in the publica- 
tion of the Annals. 

Professor T. B. Symons moved that a vote of thanks be 
extended to the authorities of the University of Minnesota for 
their kindness in offering the use of the School of Mines Building 
for the Meetings of the Society. 

On motion of Professor T. B. Symons, the Society adjourned 
to meet in joint session with the Association of Economic 
Entomologists in the afternoon. 

The Annual Public Address was given in the Handicraft 
Guild Hall at 8:00 P. M., by Professor F. L. Washburn: The 
Typhoid Fly in the Minnesota Iron Range. 

1911] Minutes of the Minneapolis Meeting 87 

Report of the Executive Committee. 

December 27, 1910. 

The Executive Committee met in the corridor of the Hotel 
Dyckman at 10 :00, with the foUowing members present : Profess- 
ors Smith, Bruner, Osborn, and Sanders. The fohowing busi- 
ness was transacted: 

List of Members Deceased during the Year 
Ending November 30, 1910. 

P. A. Hcrrick, New Brighton, Pn. G. A. West, Urbana, 111. 

G. W. Peck, Rosellc Park, N. J. Rev. J. L. Zabriskie, Brooklyn, N. Y. 

Henry Ulkp, Washington, D. C. 

The following were elected to membership in June, 1910: 
E. M. Walker, C. R. Alexander, 

Edward E. Philips, Miss A. C. Stryke. 

Alvin R. Cahn, 

The following were elected by the Executive Committee: 

Henry E. Ewing, Miss E. I. McDaniel, 

M. D. Leonard, F. H. Shoemaker, 

R. D. Whitmarsh, W. R. McConnell, 

E. W. Stafford, W. R. Thompson, 

E. O. Essig, D. Finkelstoin, 

H. R. Jennings, C. R. Plunkett, 

George G. Becker, E. W. Scott. 

The following resignations have been accepted and member- 
ship terminated: 

C. C. Adams, F. W. Powers, 

J. S. Faaborg, W. G. Wright. 

A Mares. 

The Secretary-Treasurer reported a list of eighty members, 
who, according to the rules of the Society, had been dropped for 
the nonpayment of dues. The Executive Committee referred 
this matter back to the Secretary and authorized him to write 
a personal letter to each. 

The Treasurer presented the following report of receipts 
and disbursements for the year ending November 20, 1910. 

88 ^Annals Entomological Society of America [\'ol. lY, 


Balance forward 

Received from H. Osborn, subscriptions. 

Cash received for dues, 1910 

Cash received for dues past 

Cash received for subscritpions, 1910 
Cash received for subscriptions, pasi 

H. Osborn, Xov. 11, 1910 , 



For Annals, Dec, 1909,1000 .51X2.70 

March. 1910, .SOO 192.08 

lunc, 1910, SOO 170.23 


Includes reprints, etc., clerical, typewriting 37.40 

Postage, stamped envelopes, cards. . . , 22.21 

Half-tones Annals 9.22 

Dues, notices, statements 11.50 

Express, telegrams, ledger paper, dating stamji 1.00 

Excess remittance returned to Akerlind 1 .00 

Bahmce cash on hand . - 3.52.49 


S 38.32 





Total ... .$981.04 

Of the $352.49 now on hand $1()(), the fees from Hfe members, 
is deposited in tlie Rothschild Bank of Ithaca where it is drawing 
4% interest. 

There is charged against the Society to offset dues of mem- 
bers dropped, resigned and deceased, 91 members, $239.85; 
Charges against Annals to offset subscriptions for members 
dropped, resigned and deceased, $35.00; total, $274.85. 

These charges reduce the apparent assets for the year con- 
siderably. Quite a number of these members dropped out last 
year, but the proper charge was not made on the books. 

The Secretary was instructed at the Boston Meeting to take 
a mail vote of all members and fellows of the society as to 
whether the present arrangement for separate dues and 
subscriptions to the Ann.\ls should remain in force, or whether 
they should be combined into a single fee of two dollars with 
the provisions that all should receive without further expense 
the publications of the Society. The result of this vote was as 
follows: For the amendment 182; against the amendment IS; 
blanks returned but preference not expressed 2; total 202. 
While the vote was decidedly in the affirmative, only slightly 
over one-half of the members voted. 

J. G. Sanders, Secretary. 

1011 1 M unites of the Minneapolis Meeting 89 

Report of the Committee on Nomenclature. 

The Committee has received a letter from Dr. C. W. Stiles, 
of the International Commission on Zoological Nomenclature, 
stating that it is proposed to work out the correct names of 
all the animals most intimately connected with man. In the 
course of this work, it becomes necessary to deal with the 
insect parasites of man, and it is desired that the list, as finally 
presented, shall show the correct names as determined under 
the International Code, and enumerate all the synonyms. 
Dr. Stiles suggests that this work on the insects shall be under- 
taken in the first instance by the Nomenclature Committee 
of the Entomological Society of America, in correspondence 
with the like Committee of the Association of Economic Entom- 
ologists, and such other persons as it may seem desirable to 
consult. The report so prepared should, it is suggested, be 
referred to the Committee on Nomenclature of the International 
Entomological Congress and the International Commission 
on Zoological Nomenclature, whence it would pass to the Zool- 
ogical Congress three years hence. 

Your Committee is anxious to further these plans, recog- 
nizing that the proposed list would be of great service. There 
are, however, some difficulties. The Committee of the Associa- 
tion of Economic Entomologists was formed for the purpose of 
determining the common or vernacular names of insects, and 
has not hitherto concerned itself with scientific nomenclature 
beyond printing lists of scientific names to accompany and 
define the common names proposed. Your Committee itself 
was appointed to discuss nomenclatural questions, for which 
the data were sup]50sed to be provided, and did not ex]3ect to 
have to report on matters outside of the range of nomenclature. 
It is obvious that the preparation of a complete and authentic 
list of the insect parasites of man involves many taxonomic 
questions to which nomenclature is only secondary. It is not 
understood whether the list should include only parasites in the 
restricted sense, but we suppose that in order to be of real value 
and importance, it should contain the names of various blood- 
sucking forms, Culicidae, Glossi>ia, etc., etc., which are certainly 
intimately connected with man. Taking this for granted, we 
are at once brought into contact with various difficulties, e. g., 

V)() Annals Entomological .Society of America [Vol. VI, 

those connected with the proper classification of the Culicidae, 
and under the circumstances, your committee is wholly unwill- 
ing to merely compile a catalogue from the literature, correcting 
any obvious violations of the rules of nomenclature which may 
be found. 

Probably the only way in wliicli your Committee could pre- 
pare a satisfactory' work would be through inviting specialists 
in the different groups of insects to submit their lists, which 
might be published under the signatures of their authors, and 
discussed and amended as might seem necessary. For this 
purpose mere outlines, without details, would usually suffice. 
If the cooperation of the specialists was freely given, and their 
proposals were freely discussed for a period, the Committee 
might then be in a position to bring the results together in a 
single catalogue. 

The Committee would call the attention of entomologists 
generally, to the importance of preparing lists giving the 
synonymy and indicating the generic types in their respective 
groups. Such work would go far toward permanence in 
generic designation, particularly if of such a scope as to include 
the genera of an entire faunal region, rather than accepting a 
continental or national limitation. Such contributions to 
knowledge should involve assistance from practically all workers 
in a group and your committee hopes that shortly this will 
be the general rule. 

In dealing with various matters, it is occasionally found 
that the International Code, as at present constituted, is either 
capable of more than one interpretation, or fails to settle a 
matter in dispute. We have discussed some of these questions, 
but at the present time desire only to offer the following sug- 
gestions for the consideration of the Society. It is to be under- 
stood that so far as these provisions may be different from or 
additional to those of the International Code, it is intended that, 
if they are adopted, they shall be transmitted to the Internation- 
al Committee, for consideration as amendments to the code. 

(1) Secondary homonyms, based on invalid combinations, 
shall not be recognized. This means that if a new species 
is published as A — b — , and is later wrongly transferred to 
another genus as B — b — , it is still permissible for an author 
to describe a new species as B — b — , although he may not 

1911] Minutes of the Minneapolis Meeting 91 

name one A — b — even if the species originally so named has 
been properly transferred to some other genus. 

This point is not specifically covered by the International 
Code, although the spirit of the code seems rather against it. 
It is however covered by the American Ornithologists' Union 
Code (1908 edition, p. Ivii), and correspondence shows that it 
is favored by many entomologists. 

(2) When an author describes a new species, citing several 
localities, and not mentioning any one as typical, then any 
writer following may designate any one of the localities origin- 
ally given as the type locality, provided always that nothing 
in the original name or description indicates otherwise. (If 
the name of the species has reference to any locality or to any 
collector who collected in only one of the localities cited, this 
will suffice to fix the type locality from the original publication 

The following, formulated by a member of the committee, 
is now offered for discussion without endorsement, the majority 
of the committee feeling that it requires further consideration 
or perhaps amendment. 

(3) Generic names shall not be considered as validly pub- 
lished unless the author, at the time of publication, either 
mentions an included species by its scientific (binominal) 
name, which name has been validated by a description; or cites 
a species in such a way that definite reference can be made, 
following the data given, to a previously published scientific 
name. It may be held, however, that when a genus is proposed 
with a description, and a single new species cited as type, the 
latter without description, then the generic description may 
cover both, just as if the author had given the whole combina- 
tion at the beginning, followed by "n. g. and sp.", as is fre- 
quently done. 

Differences of opinion exist as to whether the above rule, 
or the spirit of it, is in accordance with Article 25 of the Inter- 
national Code. 

H. T. Fernald, 
E. P. Felt, 



The Annals of the Entomological Society of America will be 
published by the Society quarterly and will include the Proceed- 
ings of the Annual meetings and such papers as may be selected 
by the Editorial Board. 

Papers may be submitted to any member of the Editorial 
Board and should be as nearly as possible in the form desired as 
final, preferably typewritten, and illustrations must be finished 
complete ready for reproduction. Plates must not exceed 5x7 
inches unless intended to fold. In general, papers to be accepted 
must be original, complete and previously unpublished and, ex- 
cept in connection with the proceedings, it will not be the policy 
to publish preliminary announcements or notes. Authors will 
be allowed fifty reprints gratis and additional copies at cost to 
the Society. 

Requests for information as to membership and the annual 
subscription and dues of members may be sent to the Secretary- 
Treasurer, A. D. MacGillivray, Cornell University, Ithaca, N.Y. 

Communications relating to the Annals, and all orders for 
separate copies or reprints should be addressed to the Managing 
Editor or to Annals of the Entomological Society of Amer- 
ica, Biological Building, 0. S. U., Columbus, Ohio. 


Banks, Nathan — Notes on African Myrmeleonidae . . . i 

Chamberlin, Ralph V. — The Lithobiomorpha of the 

Southeastern States 32 

Wilson, H. F. — Notes on the Synonymy of the Genera 

Included in the Tribe Lachnini 51 

Felt, E. P. — Summary of Food Habits of American 

Gall Midges. 55 

Jensen, J. P. — The Structure and Systematic Import- 
ance of the Spermatophores of Crickets 63 

Herrick, Glenn W. — Notes on the Life History of 

the Larch Case-Bearer (Coleophora Laricella.) 68 

GiRAULT, A. A. and Zetek, James — Further Biologi- 
cal Notes on the Colorado Potato Beetle, Leptino- 
tarsa lo-lineata (Say) 71 

Sanders, J. G. — Minutes of the Minneapolis Meeting, 84 

The regular annual subscription price for the Annals is, in 
the United States, Cuba, Porto Rico, Hawaii and Mexico, $3.00; 
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orders should be drawn payable to Annals Entomological 
SoaETY OF America, and addressed to Biological Building, 
O, S. U., Columbus, Ohio, U. S. A. 

Volume IV. Number 2. 


The Entomological Society of America 

JUNE, 1911 



ITHACA, N. Y. Washington, D. C. 


G0EI.PH, Ontario, Canada. Boston, Mass. 


Boston, Mass. PnitADEtPHiA, Pa. 


Stanford Uni\'., Cai.. Urbana, Ili.3. 

HERBERT OSBORN, Managing Editor, 
Columbus, Ohio. 


Elnlered as second class master April II, 1908, at the Post Office at Columbus, Ohio, 
under the Act of Cohgtess ot March 3, 1879. 

The Entomological Society of America. 

FOUNVED 190e. 

OFFICERS -1911. 

Pre jiieni— Herbert Osborn Columbus, Ohio 

First Vice-President — Lawrence Bruner Lincoln, Nebraska 

Second Vice-President — A. D. MacGillivray. Ithaca, New York 

Secretary-Treasurer — A. D. MacGe-LIVR-W Ithaca, New York 

Executive Committee— Tbe Officers, and J. H. Comstock, W. M. Wheeler, 
J. B. Smith, C. J. S. Bethune, Henry Skinner, A. D. Hopkins. 
Committee on Nomenclature— R. T. Fernald, E. P. Felt, T. D. A. Cockerell. 

Price List of Publications. 

Annals, Vols. I, II and III, complete, each $3.00 

Annals, Vols. I and II, Parts 1, 2 and 4, each 1.00 

Annals, Vols. I and II, Part 3, each 50 


Comstock, J. H.— A Note on the Habits of the Wall-bee Chalicodoma Muraria .10 
Petrunkevitch, a. — Contributions to Our Knowledge of the Anatomy and 

Relationships of Spiders 15 

Girault, a. Arsene — A Monographic Catalogue of the Mymarid Genus 

Camptoptera Foerster, with Description of One New North America 

Form - 15 

Davis, John J. — Studies on Aphididae II 20 

Hilton, Willwm A. — The Tracheal Supply in the Central Nervous System of 

the Larva of Corydalis Co/nuta .- .25 

Nelson, Jas. A.^Evolution and Adaption in the Palpus of Male Spiders 15 

Webster, F. M. — Investigations of Toxoptera Graminum and Its Parasites . . .25 

Hayhurst, Pal^— Observations on a Gall Aphid (Aphis Atriplicis L.) 15 

Patch, Edith M.— Homologies of the Wing Veins of the Aphididae Psyllidae, 

Aleurodidae, and Coccidae 50 

Hine, James S. — Robberflies of the Genus Asilus 50 

Chamberlin, Ralph V. — Some Records of North American Geophilidae 

and Lithobiidae, with Description of New Species 25 

Davis, John J. — Two New Genera and Species of Aphididae 10 

Poltlton, Prof. E. B, — Mimicry in the Butterflies of North America 60 

Townsend, Chas. H. T. — Descriptions of New Genera and Species of 

Tachinidae 10 

Cockerell, T. D. A.— Fossil Insects from Florissant 10 

McGiLLiVRAY, A. D. — A Synopsis of the North American Species of Scoli- 

oneurinae 20 

Hambleton. J. C— Life History of Coizus Lateralis Say 10 

For Reprints from Volume I, see preceding Number. 


Biological Building, O. S. U., Columbus, Ohio. 



The Entomological Society of America 

Volume IV J U N E , 1 9 1 1 Number 2 


By Anna H. Morgan. 
Limnological Department, Cornell University. 

The following paper is a preliminary study of the ecology 
of the May-flies in the streams about Ithaca, N. Y., more 
especially in Fall Creek. In these, as in most fresh water streams, 
the nymphs of this order are abundant. In Fall Creek they 
are the dominant insects of the stream during the months of 
^."^pril, May and June, and by their fine adaptations to diverse 
environments they offer a satisfying field of study to any brook 
traveler. The nymphs may be easily secured, but only imagoes 
exist in most collections, and these usually as dried distorted 
specimens whose life-histories are little known. The winged 
or aerial life lasts but a few days at most; the nymphal or 
aquatic life may extend over two or three years. The imago 
exhibits great specialization of parts concerned with reproduct- 
ion and more striking atrophy of other parts than may be seen 
any where else among insects. Imagoes of all the groups are 
remarkably alike in superficial appearance. The nymphs, on 
the other, hand, display a series of adaptations as diverse as 
their environments. Only by rearing specimens from nymphal 
to adult life may these two stages be linked together. Many of 
the life-histories of those species found in Eastern North 
America have been made known. It has been with the hope 
of adding to the number of these life-histories, as well as with 
the purpose of gaining more knowledge of the habits of those 
already known, that this study has been begun. The earlier 
American workers. Say, Hagen and Walsh* scarcely took up 

* Walsh, B. D. On the pupa o£ the Ephemerinous genus Baetisca Walsh. 
Proceed, of the Ent. Soc. of Philadelphia. 1864. pp. 200-206. 

* Walsh, D. B. List of the Pseudoneuroptera of Illinois. Proceedings of the 
Natural Sciences of Philadelphia. 1862. 


94 Annals Entomological Society of America [Vol. IV, 

the rearing of nymphs but by their descriptive work they laid 
the foundation for the life-history studies which have fol- 
lowed. Berryt '03 reared and described the nymphs of Habro- 
phlebia americana Banks, (which nymph is not a Habrophlebia 
but a typical Leptophlebia), Blasturus cupidus Say, and Calli- 
baetis ferruginea Walsh. A note on the nymphs of the genus 
Tricorythus was published by Cockerell and Gill '06{. The 
largest number of life-histories of Eastern North American 
forms has been written by Professor James G. Needham in 
Bulletins 47, 68, and 86 of the New York State Museum, and 
includes the following species: 

Bull. 47. Bull. 86. 

Heptagenia pulchella Walsh. Chirotonetes albomanicatus Needham- 

Baetis pygmea Hagen. Ameletus ludens Needham. 

Siphlurus altematus Say. Choroterpes basalis Banks. 

Caenis diminuta Walker. Baetis pygmea Hagen. 

Hexagenia variabilis Eaton. Callibaetis skokiana Needham. 

Ephemera varia Eaton. Ephemerclla bispina Needham. , 

Bull. 124. Caenis allecta Needham. 

Ephemerella dorothea Needham. Leptophlebia praepedita Eaton. 

Potamanthus diaphanus Needham. Heptagenia interpunctata Say. 

Ecdyurus maculipermis Walsh. 

By Mr. W. E. Howard. 
Polymitarcys albus Say. 

With the exception of Callibaetis skokiana, Ephemerella 
bispina, Ephemerella dorothea, and Potamanthus diaphanus, 
all of these species have been taken in or near Fall Creek. 
For some of these further biological data have been secured. 
In addition to them eight species have been bred which it is 
believed have not been before recorded. These are all from 
Fall Creek with the exception of one, Ephemerella cornuta, 
reared for me by Miss Lucy W. Smith at Salisbury, Connecticut, 
and here included in the Ephemerella group. The life-histories 
which are given are those of Iron fragilis, sp. nov., Epeorus 
humeralis, sp. nov., Ephemerella rotunda, sp. nov., E. tuber- 
culata, sp. nov., E. cornuta, sp. nov., E. deficiens, sp. nov., 
E. plumosa, sp. nov., and E. spinosa, sp. nov. The descrip- 
tion of the female imago has been added to Prof. Needham's 
life-history of Ameletus ludens, and the Caenis allecta which he 
placed provisionally in that genus has on rearing been estab- 
lished in Tricorythus. 

t Berry, Edward. New or Hitherto Unknown Ephemerid Nymphs of the 
Eastern U. S. Am. Natural. Vol. XXXVII, pp. 25-31. 1903. 

X Cockerell, T. D. A., and Marie Gill. Tricorythus, a Genus of Mav-flies. 
Univ. of Colo. Studies, Vol. III. No. 3, 1906. 

1911] May-Flies of Fall Creek, N. Y. 95 


The vicinity of Ithaca consists of two highlands between 
which hes the basin of Cayuga Lake. The west highland 
known locally as West Hill is a long regular slope, while East 
Hill upon a terrace of which Cornell University stands, is fur- 
rowed with gorges made by streams flowing downward to the 
lake. On the gradual incline of the Eastern highland these 
streams flow along as quiet meadow brooks, or broadening out 
over stony beds are caught in a maze of ripply shallows, but on 
reaching the steep terraces of the highland they plunge down- 
ward through the narrow gorges by a succession of cascades 
till they come to the plain below. These streams coming far 
from their source and fed by many tributary waters are flooded 
and turbulent in the sjDring, but gradually dwindle to mere 
brooklets with trickling falls during the mid and late summer, 
when the tributaries fail of their supply. Few of the main 
streams become wholly dry. In March and April rich flora 
and fauna spring from their banks and waters, while through 
the dry season they supply enough water for the maintenance 
of life and the reproduction of another generation. Fall Creek, 
which bounds the Cornell Campus on the north, is a type of 
these streams. 

About one mile east of the campus Fall Creek flows over a 
broad nearly level bed thickly strewn with flat stones and rocks 
which project from the water except at periods when the stream 
is swollen. On one side the creek is bordered by a soft sandy 
shore, on the other by a shelving ledge. Beyond this point, 
where the ledge gives place to soft drift, there is a series of 
permanent pools which mark the entrance of a small tributary 
spring. A cross section of this upper portion of the stream 
represents a variety of situations great enough to shelter widely 
different types of May-fly nymphs. Clinging to the surfaces 
of the stones in the mid current are the flat nymphs of Epeorus, 
Iron, Ecdyurus and Heptagenia; clambering in the trash which 
has collected between the stones are the nymphs of Ephemerella; 
on the sheltered surfaces or in the quiet border waters are Hep- 
tagenias about to transform in company with Leptophlebia, 
Siphlurus, and Ameletus; hidden in the sandy sweeps are Caenis 
and Tricorythus and burrowing in the soft muck banks are 
Hexagenia and Ephemera. Changing from this gradual descent 

96 Annals Entomological Society of America [Vol. IV, 

Fall Creek cuts downward through a narrow gorge, widens into 
the artificial pond known as Beebe Lake, hurries through a 
deep gorge and over a series of falls, cascades and riffles to the 
marsh below. This lower creek is inhabited by those true 
dwellers of the rapids, Chirotonetes albomanicatus and Baetis 

To the north of the lower portion of Fall Creek a small 
streamlet known as Pleasant Brook follows a parallel course to 
the lowlands. Its pools and cascades shelter a fauna similar 
to that of Fall Creek if somewhat less rich. This brook 
possesses the advantage of small size which makes its study easy. 
Devoid of tributaries to flood it in time of rain, and shut in by 
shrubbery, this stream furnished a safe place for the rearing 
cages of nymphs which were captured in Fall Creek. 


Rearing and collecting were begun on April 1, and continued 
to August, 1, after which only irregular collecting trips were 
made to the Creek. The only satisfactory method of rearing 
May-flies is one which keeps them in their own environment 
or in conditions closely imitating it. For this purpose Prof. 
Needham used a cylindrical cage made of wire cloth with a 
cheese-cloth cover. Such cages are the most convenient for 
carrying in a knapsack and many May-flies have been success- 
fully reared in them. The space within them, however, is 
small and all surfaces are perpendicular to the water. If the 
imago becomes entangled, or if it is not strong enough to keep 
its footing on the upright surface it falls back into the water 
and drowns, or at least will never be able to transform. When 
two or three insects are in the same cage, particularly if the cage 
is in a strong light, there is danger of one or all falling into the 
water. For these reasons I have designed another cage, which 
though less conveniently carried about, has the merit of being 
more roomy and of supplying one slanting surface. This cage 
may be made of fine copper or galvanized wire cloth. A stiff 
cloth which will not bend easily will make the best cage. The 
cages which I used in Fall Creek were about five inches square 
on the bottom and five inches in height. Such a cage is easily 
cut and folded from a single piece of wire cloth. In the dia- 
gram shown in Fig. 2, the continuous heavy lines represent the 
cut edges, the lighter lines the folded edges of the laps, and the 


May- Flies of Fall Creek, N. Y. 


dotted lines the angles of the bottom, back, sides, front and 
cover. The laps on the sides should be folded over the cut 
edges of the bottom and the front and then securely fastened 
with solder. The cover may then be pushed down and secured 
by a wire catch or by a rubber band placed about the cage. 
When in use a stone should be placed in the bottom of the cage. 
This will serve the double purpose of keeping the cage upright 
and of providing a foot hold for the nymphs enclosed within it. 
In Fig. 1 the completed cage is shown inserted in the water. 
A represents the stone placed in the cage. 

Fig. I. Cage for rearing May flies, showing 

position in the water. 
Fig. 2. Diagram to show construction of cage. 

It is best not to insert the cage much more than two inches 
in the water except where a lowering in the stream is expected. 
Nymphs confined in this cage will naturally crawl up the slop- 
ing side for emergence and the sub-imagoes will find an easy 
grade on which to walk up to the light. The sub-imagoes will 
sit on the under side of the cover, but if it be lifted with some 
care the insects may be safely transferred from the cage to the 
collecting box. 

98 Annals Entomological Society of America [Vol. IV, 

Many times nymphs are collected for rearing in places not 
often visited. Such nymphs may be transported alive in jars 
of their native water with plants or stones to furnish forage and 
foot-hold. Fragile forms like Epeorus and Heptagenia may be 
better carried in a can of cotton or sphagnum thoroughly sat- 
urated with water. The nymphs should be carefully placed on 
the surface with a thin, very wet layer above them. If carried 
in this way they will arrive at their destination in much more 
perfect condition than if jostled about in a can of water. If a 
running water aquarium, or better, a convenient small stream 
is not available, the nymphs brought home for rearing may be 
placed in flower-jDot saucers in which rapid evaporation will 
keep the water sweet. Small stones projecting from the water 
should be provided for emerging places. A cylinder of wire 
cloth with cheese-cloth top may be slipped over the dish so 
that the sub-imago may be easily caught. 

In large aquaria where several kinds of insects are kept, 
care should be taken to exclude carnivorous beetles, and 
dragon-fly and damsel-fly nymphs for which May-fiies are 
choice food. May-fly nymphs are mostly herbivorous and need 
only a supply of diatom-covered stones for forage and some 
aquatic plants like Chara or Nitella upon which they may depend 
for foot-hold and hiding. Needless to say the temperature of 
such an aquarium should be kept as nearly as possible to that 
of the streams. Of the nymphs which I have endeavored to 
keep in the aquarium of Cornell University, Blasturus cupidus, 
Callibaetis fluctuans, and Siphlurus alternatus proved most 
hardy. These lived from one to four weeks in rather adverse 
conditions, the water in the aquarium having been treated 
with aluminum sulphate for drinking purposes. 


The May-flies found during the past summer in or very near 
the Creek will be grouped under the three family heads given 
by Prof. Needham in Bull. 86. Notes and description of new 
species are given under their respective headings. 


Polymitarcys albus Say. 

Two sub-imagoes were captured near night-fall on June 20. 

1911] May-Flies of Fall Creek, N. Y. 99 

Potamanthus sp. 

Only two partly grown nymphs of this form were found. 
Both were taken on June 29 in sandy mud washed by a gentle 

Hexagenia variabilis Eaton. 

Full grown nymphs of Hexagenia variabilis Eaton the 
largest of our burrowing May-flies, were found abundantly 
on the sunny afternoon of May 16 in company with Ephe- 
merella and the large dragon-fly Cordulegaster. At the 
place where they were Fall Creek is a leisurely brook 
meandering through sparse woodland and open meadow, 
and hemmed in by soft muck banks. In one such area 
the sloping banks were mined by Hexagenia nymphs, the 
open burrows showing only two or three inches apart. 
Most of the burrows were apparent by their round open- 
ings, but from some hairy caudal setae protruded at full 
length. When a nymph was pulled out it speedily began to 
burrow again, placing the forelegs together with the blade-like 
tarsi held verticahy. It next pressed them forward and outward 
at the same time wedging the head between them in the cavity 
thus made. This movement was followed by a sudden lurch 
of the body forward accompanied by wriggling of the abdomen. 
During these motions the second pair of legs was folded close 
up to the body, while the third pair was held outstretched 
ready to brace against the mud. These motions rapidly 
repeated enabled the nymph to bury itself in a surprisingly 
short time. Some of the soft ooze taken from where the bur- 
rows were most numerous was later examined in the laboratory 
and found to be packed with diatoms. Stomachs of two of the 
nymphs were found full of silt and diatoms showing that the 
nymphs had found plentiful forage as they burrowed. Between 
two and five o'clock of this afternoon about twenty-five sub- 
imagoes emerged within a few yards distance. They flew up 
slowly and usually settled on low shrubs. Many were cap- 
tured on near-by alders two to five feet from the ground where 
the yellow markings on their bodies and wings made them 

100 Annals Entomological Society of America [Vol. IV, 

Ephemera simulans Walker. 

No representatives of this genus were collected previous to 
June 14 when a female specimen was captured at large near 
upper Fall Creek. From June 23 to 30 full grown nymphs 
were found about two inches below the surface in a muddy 
basin connected with the main stream. Near this place a 
swarm of Ephemera consisting of three to four hundred indi- 
viduals was dancing about fourteen feet in air at half past 
seven o'clock in the evening of June 29th. Their steady rising 
and falling continued over the same area as long as the light 
kept them visible. One female captured from the swarm was 
placed on the surface of water in the laboratory. She was 
unable to fly and lay prone upon the surface. Immediately 
the last four segments of the abdomen began to move spas- 
modically and eggs poured forth from the oviducts. At the 
end of one minute the abdomen was empty, and the glass 
spread with a single layer of white, firmly adherent eggs, 
easily distinguished with the naked eye. 

No Ephemera nymphs were found in the lower Creek up 
to this time, that cleaner portion being nearly devoid of mud. 
On the first of July, however, the water in Beebe Lake was 
allowed to run off, bringing into the Lower Creek large quanti- 
ties of mud. Three days later the shores below the dam were 
again examined. Tracks similar to those made by earth-worms 
covered the bottom near the shore-line. Nymphs were crawl- 
ing over the surface and setae could be seen projecting from 
many burrows. From an area of about ten square feet thirty 
nymphs were removed. 


Represented entirely by dwellers in rapid water,, this family 
is the dominant one in number, and the most homogeneous in 
nymphal form. It is represented here by four of its six North 
American genera Heptagenia, Epeorus, Ecdyurus, and Iron, 
given in order of their relative abundance. As a family the 
Heptageninae has taken possession of the rapid, thoroughh' 
oxygenated water and the alga-covered stones of the middle 
stream, apparently coming into the calmer waters only at 
transformation time. In order to secure and keep this posi- 
tion against the mechanical force of the water acting upon them 

1911] May -Flies of Fall Creek, N. Y. 101 

all alike, they have been forced into a series of similar adapta- 
tions. The principle feature of these adaptations are: a gen- 
erally depressed body; dorsally placed eyes and flaring margins 
to the head; spreading legs with flattened femora and lateral 
pectinations on the claws; a series of over-lapping gill lamellae, 
and fiat widely diverging caudal setae. 

Heptagenia interpunctata Say. 

Five species of the genus Heptagenia were collected and 
reared during the summer, but only the very common H. inter- 
punctata will be listed until further work can be done upon 
them. H. interpunctata was common from April 30, when I 
found nearly grown nymphs beneath the fiat stones in a trib- 
utary of the creek up to August 1, after which little collecting 
was done. During this time many emergings were observed, 
the greater number occurring between two and five o'clock in 
the afternoon. 

Ecdyurus maculipennis Walsh. 

Associated with Heptagenia and Epeorus, but with a pref- 
erence for more gently flowing water, Ecdyurus maculipennis 
is a frequent dweller in the quieter border water of swift cur- 
rents. My first collections are dated June 3. From this date 
till July 2,5 it was a common associate of Heptagenia and 
Epeorus, slightly smaller and a swifter runner than either of them. 
It also bears transportation and change of water with greater 

Iron fragilis. 

The nymph of this species was described by Prof. Needham 
in 1905. In addition to this description there have been but 
two records of this genus in North America, Iron nitidus,* 
Oreg. Cal. and I. longimanus,t Colorado. The first nymphs of 
this species collected during the summer were found on May 9, 
in Coy Glen, the stream from which Prof. Needham's speci- 
mens were taken. Here thick growths of Cladophora and 
diatoms support an abundant May-fly population. The 
nymphs live in the swiftest water, on the under side of the stones, 
in the falls, or on the smooth rock floor. A census of the inhab- 

* Iron nitidus Eaton. Rev. Monog. 246, 1885. Oreg. Cal. Banks, Cat. Neur. 
Insects. Am. Ent. Soc. '07. 

t I. longimanus, Eaton, Ent. Mo. Mag. XVIII, 26, 1881. Rev. Monog. 245, 

102 Annals Entomological Society of America [Vol. VI, 

itants of twenty stones measuring about seven by eleven inches 
revealed the following inhabitants listed in the order of abun- 
dance, Simulium, Blepharocera, May-flies (Epeorus, Iron. 
Baetis), Stone-flies and Parnidae. In competition with such 
structures as the sucking disks of Blepharocera and the limpet 
like form of the water-penny (Parnidae), Iron has developed 
a successful hold fast of its own. The first pair of gill lamellae 
are very large and scoop shaped with their hinder edges over- 
lapping the succeeding lamellae, and their front edges meeting 
beneath the posterior portion of the thorax. The lamellae 
diminish in width posteriorly and the last pair are incurved 
beneath the abdomen. The edges of the lamellae have a thick- 
ened border and when closely pressed to the supporting surface 
a successful holdfast is formed. This is one of three closely 
allied genera, Epeorus, Iron, and Rhithrogena, which represent 
remarkable modifications for life in rushing water. All three 
possess closely overlapping gill lamellae and but two caudal 
setae. The main differences are in the shape of the ventral 
abdominal disk which in Epeorus is incomplete, the first and 
last pairs of lamellae being distant ; in Iron nearly complete, the 
first and last pairs of lamellae nearly meeting; in Rhithrogena 
the disk is completed by the perfect apposition of these lamellae. 
The mouth-parts (PI. X, fig. 1) are completely hidden from 
above by the flaring margin of the head with its bordering fringe 
of soft hairs. Viewed from beneath the small labrum (PI. 
X, fig. Ir) may be seen curving downward and backward over 
the tips of the mandibles and maxillae (md. a. and mx. a) to 
meet the median flaps of the broad labium (1. 1. e.) The labium 
is flat and its outer surface (1) fits close down to the surface upon 
which the insect is foraging. Along the anterior margin of the 
outer surface of the labial palp is a series of incurving hairs 
(1. 1. a.), behind these a set of overlapping plates, and still 
farther backward a single strongly chitinized scraper, (1. c.) 
On the inner surface of the palp is a semi-circular patch of 
inwardly directed hairs (1. b.) Closely apposed to the inner 
surface of the labium is the outer surface of the hypopharynx 
(fig. 1 hy), made up of two lateral, and one median portion, 
distinct, except at their bases. The separation of the two 
lateral portions from the median portion leaves a gutter-like 
trough between them on the inner side. The labial palpi are 
freely movable horizontally. They are moved outward, then 

1911] May-Flies of Fall Creek, N. Y. 103 

pressed slightly downward and inward when feeding. Some- 
times the labial palp is slipped between the median (fig. 1, 1. e.) 
and the lateral flaps (fig. 1, 1. d.), sometimes over the inner 
sides of both. In the first case the plates on the outer sides of 
the palp are brushed by the hairs on the inner side of the median 
flap while the semi-circular patch of hairs (1. b.) on the inner 
side of the palp is brushed by those on the outer side of the lateral 
palp. In the second and more frequent case the outer side of 
the palp is brushed by both flaps and the semi-circular patch 
of hairs fits into the chitinized gutter on the adjacent surface 
of the hypopharynx (hy. a.) Directly above the median por- 
tion of the hypopharynx are the grinding surfaces of the maxillae 
(mx. b.) with those of the mandibles (md. b.) directly above 
them. Food scraped inward by the labial palps is evidently 
deposited beneath the hypopharynx. From here it is probably 
sucked up into the mouth cavity through the slits between the 
median and lateral folds. The maxillary palps act in a manner 
similar to those of the labium, but because of their position 
above the hypopharynx, they must be able to place the food 
directly in the mouth cavity. The inwardly curving hairs on 
the lateral borders of the labrum help to keep the food in the 
mouth while it is being chewed up. 

The foregoing observations were made by placing a nymph 
in a shallow dish of water, throwing a strong light upon it and 
studying it with a binocular microscope. The nymph was 
uneasy in this unnatural position and kept its mouth parts 
continually moving. Nymphs may be studied tmder more 
natural conditions if they are placed in a glass-bottomed box 
with a small amount of forage. The box may be placed upon 
some support which will elevate it above the table. The bot- 
tom may then be tilted at any easy angle and the nymphs 
studied with a hand lens from beneath. 

Occurrence, habitat. The dates on which nymphs or ima- 
goes were collected range from May 1 to June 15. These 
nymphs were all taken in cool, shaded waters and were most 
abundant during the early part of the season. The data for 
those reared in cages is as follows : 

1 male emerged May 11, a. m., transformed May 13, a. m. Coy Glen. 
1 male emerged May 12, a. m., transformed May 14, a. m.. Pleasant Brook. 
.3 females emerged May 30, p. m., transformed June 1, p. m. Pleasant Brook. 
1 female, 1 male emerged June 10, a. m., transformed June 12, M. Fall Creek. 

1U4 Annals Entomological Society of America [Vol. IV, 

Between the hours of two and four of June 15 about twenty 
emergings were witnessed from one view-point in a narrow swift 
portion of Coy Glen Brook. The nymphs popped from the 
surface of the water and flew unsteadily upward in the sunlight 
for about 20 or 30 feet when they veered into the tree tops or 
settled on the side of the gorge. 

Iron fragilis sp. nov. 

Length of body Length of setae Length of Wing 
Male imago 7 m. m. 21 m. m. 7 m. m. 

Male subimago 6.5 m. m. 14 m. m. 

Female imago 7 m. m. 15 m. m. 

Female subimago 6.05 m. m. 14 m. m. 

Male imago. (Plate X, figs. 2, 3 and 4). (In alcohol). Body 
extremely delicate and fragile. General hue dull yellowish white, 
appearing hyaline in segments 2-7 of the abdomen. Eyes, conspicuous, 
grayish. Head, parchment color with the eyes gra^-, darker below; 
antennae, light brown except for white basal segment; ocelli, ringed with 
conspicuous, broad band of dark brown. Thorax, yellowish shading 
brownish above, pale below. Pronotum deeply notched behind. The 
lateral lobes of the pronotum and the median portion of the meso- 
thorax shaded brown. A sub-triangular area of brown on either side 
of the median posterior elevations of mesothorax and metathorax. 
Legs, dull yellowish, the femora with a conspicuous dark brown spot 
at the middle. Tarsi with joinings and claws brown. Forelegs (PI. 
X, fig. 3) longer than the body, its tarsal claws identical with those of 
the other two. Wings, (PI. X, fig. 2) hyaline, sub-hyaline near the tip 
in costal and sub-costal regions. Abdomen. Tergites 1-9 with their 
posterior borders delicately shaded by transverse bands of brown growing 
more distinct posteriorly. Stemites, pale. Forceps pale, broad at the 
base and conspicuous (PI. X, fig. 4). Setae 2, pale slightly brownish 
at the base. 

Male sub-imago. The sub-imago differs most markedly from the 
imago in the following respects. The forelegs and setae are shorter, 
the forceps less conspicuous, and the wings are of the usual grayish 
sub-hyaline appearance. Head with occiput brownish. Bands of 
ocelli "less prominent than in the imago. Thorax, brown, prominent 
ridges of mesothorax and metathorax shaded with brown, but without 
defaiite sub-trangular markings. Pleurae and sternum with ridges 
shaded brown. Coxae suffused with brown. Abdomen with tergites 
1-10 suffused with brown, the trans\'erse band of brown more distinct 
than in the imago. An irregularly shaped patch of white in the center 
of each segment near the pleura. The posterior lateral angles of the 
tergites whitish. 

Female imago. Body heavier than that of the male, all over didl 
yellowish color; legs of nearly equal length; setae 2. Head with eyes 

1911] May-Flies of Fall Creek, N. Y. 105 

distant and dark grayish, a whitish area of the head showing between 
them. Thorax, with the pronotum distinctly hned with brown. Sub- 
triangular areas on mcsothorax and metathorax present but less dis- 
tinct than in the male. Abdomen with tergites shaded with brownish, 
a distinct transverse band of brown near the posterior border. Stemite 
7 produced backward in a rounded lobe whose posterior edge touches 
that of stemite S. Stemite 9 produced backward in a lobe with a shal- 
low median indentation on its posterior margin. 

Female sub-imago. General color much darker than the male 
imago. Thorax shaded all over with brown. Stemite 7 produced 
backward only half the length of stemite S. The prolongation of ster- 
nite 9 much less pronounced and its posterior margin barely indented. 

Epeorus humeralis. 

(PI. IX, fig. 1, 2, 7.) The genus Epeorus is represented in 
Fall Creek by this single species. It is closely allied to Iron 
but dififers from it in the greater distance between the lamellae 
of the first and last pairs of gills. These nymphs are much 
larger than those of Iron fragilis, but are harder to transport 
because of the extreme brittleness of the bases of the lamellae. 

Occurrence, habitat. Full grown nymphs and imagoes were 
taken at various dates from May 25 to July 15. This species 
is very common in the swift waters associated with Heptagenia 
and Baetis. 

Epeorus humeralis sp. nov. 


Length of body Length of setae Antennae 

Male imago 10 m. m. 20.5 m. m. 

Male subimago 9 " 12.5 

Female imago 9.5 " 13 

Female subimago 10 " 11.5 

Nymph 11 " 11 " 2.5 m. m. 

Male imago. (PI. IX, fig. 2, 7). (Live specimen). General color 
dull yellowish, becoming sub-hyaline on segments 2-5 of the abdomen. 
Conspicuous dark brown spots at middle of femora in all winged stages, 
larger than those in Iron fragilis; the body less fragile than that of the 
preceding species; the humeral cross vein blackish; the foreleg about 
three quarters the length of the body. Head, yellowish white; eyes 
conspicuous olive green, intersected in the lower portion by a brown 
band, ventral edge margined by a narrow black band border behind, 
this in turn by a white band of the same proportions; antennae brown, 
a brown ridge extending from their basal segments to the inner margins 
of the eyes; carina on middle of front brown; ocelli ringed with olive. 
Thorax, above yellowish white, translucent; pronotum partially hidden 
by the eyes, its lateral lobes shaded with dark brown; mesonotum 
buffy with edges brown and elevations lighter; scutellum fuscous. 

106 Annals Entomological Society of America [Vol. IV, 

Below, pale whitish yellow, mesonotum mar^ned at the apex with a 
transverse band of brown. Pleurae and coxal areas pale ^\-ith irregular 
shadings of dark brown. Legs whitish; coxae with dark brown spot; 
femora with conspicuous brown spot and a brownish band just before 
the apex; more distinct on the anterior legs; tarsi with flap and claw 
similar to that of Iron fragilis (PI. X, fig. 3) and identical in all three 
legs. Wings hyaline with the humeral cross vein blacldsh brown (in all 
adult stages). Abdomen, pale whitish, translucent on segments 2-5. 
Tergites with distinct transverse brown pencilings on their posterior 
borders and a median broken line of brown more distinct on the posterior 
portions of the tergites. Stemites pale without markings. Forceps 
pale, (PI. IX, fig. 2). Setae whitish. ' ' 

Female sub-imago. General color of the body slightly darker than 
that of the imago. Abdomen darker, less translucent. Wings grayish, 
sub-hyaline. Thorax with markings less sharply defined and general 
color darker than that of the imago. Abdomen with the tergites suf- 
fused with browTush. 

Female imago. (Live specimen). Forelegs shorter than those of 
the male. In specimens containing eggs the abdomen is a bright salmon 
pink, which does not fade readih' in alcohol. Head, with eyes similar 
to those of the male, but smaller and distant being separated by a broad 
unmarked portion of the occiput. The dark bands of the ocelli are in- 
complete and a trifle narrower than those of the male. Thorax, with 
the pronotum exposed showing the prominent median indentation of its 
posterior margin; an area on either side the median line with a longitudi- 
nal blotch of brown. Abdomen with the tergites slightly darker brown 
than in the male. Stemite 7 prolonged two thirds the length of stemite 
8. Stemite 9 slightly shorter and but shallowly notched on its posterior 

Nymph. (PI. IX, fig. 1). Body depressed, widest across the meso- 
thorax gradually tapering to the last abdominal segment. Nymph 
larger and broader than Iron fragilis with its lamellae flaring; the lamel- 
lae of the first and last pairs of gills distant from one another and the 
tracheation of the lamellae conspicuous. Posterior lateral angles of the 
abdominal seginents produced into backwardly directed spines which 
guard the bases of the lamellae (PI. IX, fig. 1). Color, olive-greenish 
blotched and shaded with brown, pale below. Head with eyes and 
ocelU prominent, the latter with dark lunate bands on their inner margins. 
Antennae slender and bare. Lateral margins of the pro thorax rounded 
anteriorly, and flaring. Legs flattened with a row of rather long 
soft hairs on the posterior margins of the femora and tibiae. General 
shape conical, rounded above, slightly flattened below. Gill lamellae 
obliquely reclinate with fasciculate filaments on the dorsal side of their 
bases. Setae 2, about as long as the body. 

This heterogenous group was represented in the summer's 
collecting by Blasturus cupidus, Leptophlebia praepedita, 
Leptophlebia mollis, Choroterpes basilis, Callibaetis fluctuans, 

1911], May-Flies of Fall Creek, N. Y. 107 

Ephemerella excrucians, Caenis hilaris, Siphlurus alternatus, 
Chirotonetes albomanicatus, Ameletus ludens, Tricorythus 
allectus, Ephemerella rotunda, sp. nov., E. tuberculata, sp. 
nov., E. cornuta, sp. nov., E. deficiens, sp. nov., E. plumosa, 
sp. nov., and E. spinosa, sp. nov. All of these were taken in 
Fall Creek except the before mentioned E. cornuta. The 
descriptions of these species of Ephemerella is here given 
together with that of the female imago which has been added 
to Prof. Needham's life-history of Ameletus ludens. Tricory- 
thus allectus, which was placed in that genus by Prof. 
Cockerell has been established there by several rearings. 

Blasturus cupidus Say. 
From April 20 to May 1 this species was the dominant May- 
fly of the quiet pools. On April .30 observations were made in 
a sheltered pool, tributary to Fall Creek. This pool was about 
fourteen feet long by five feet wide, carpeted with decaying 
leaves, and bordered on one side by a thick mat of spirogyra. 
Between 11:30 and 12:30 o'clock in the bright sunlight, about 
forty nymphs emerged and the sub-imagoes were captured. 
A few records of individual emergings were taken of which the 
following is typical. A nymph appeared from beneath some 
leaves, came close to the surface of the water and swam about 
there till a stick was found which projected out of it. It imme- 
diately clambered up the stick, thrust its head out into the air 
and rested there with its gills motionless, but apparently swal- 
lowed large gulps of air. Very soon a median split appeared 
in the mesothorax, widened toward the prothorax and then to 
the metathorax. The head and eyes of the sub-imago appeared, 
the mesothorax, then the metathorax, and finally by a sus- 
stained pull, terminating in a jerk, the wings were extricated 
from the wing pads and erected. Apparently exhausted by this 
effort the insect then paused with the posterior portion of the 
abdomen and the setae still lying loosely in the cast skin. By 
another jerk the body was wholly freed from the skin. The 
insect rested an instant upon the water's surface with its setae 
held widely divergent and upward, in their natural alert posi- 
tion. Immediately after this it fluttered upward and settled 
on a low shrub. The entire time from the appearance of the 
nymph to the completion of its emerging was ten minutes. Of 
this period, one minute was occupied in swimming, one in taking 
in air, and three minutes occurred between the appearance of 

108 Annals Entomological Society of America [Vol. IV, 

the median slit in the nymphal skin and the complete freeing 
of the sub-imago. After its first short flight the insect remained 
on the shrub for about five minutes before disappearing higher 
up in the air. This custom of resting upon near-by objects is 
a habit varying with the species and apparently also with weather 
conditions. For example, sub-imagoes of Chirotonetes albo- 
manicatus which usually fly upward immediately on emerging 
in clear weather walk about for a while on the shore, or take 
very short first flights on dark days. The period of greatest 
effort during emergence is that which precedes the splitting of 
the nymphal skin. The splitting is doubtless urged on by the 
distention of the alimentary canal which is caused by the air 
or water which has been swallowed. 

Mating flights of Blasturus cupidus were observed over 
Fall Creek in the late afternoon of April 31, the height of their 
transformation season. About thirty individuals flying in 
close ranks rose and fell at varying altitudes of ten to thirty 
feet. When at their greatest height they were scarcely dis- 
tinguishable against the sky, but when they were lowest the 
forelegs and the setae might be discerned. The forelegs were 
held stiffly, straight forward from the head, and the setae, 
projected at a wide angle behind, appearing to vibrate as the 
insects swung downward. After a few moments of ecstatic 
rising and falling, one of the individuals flying high in the 
swarm descended to one of the lowest, coupled with it and 
veered obliquely downward and across the stream. When 
about to alight on the opposite shore the two separated, one 
disappeared and the other turned back and flew close to the 
surface of the stream frequently brushing the water with the 
abdomen. Such matings were three times observed. At- 
tempts to capture a fertilized female failed. The eggs brushed 
from the abdomen into the rapidly running water were, of 
course, impossible to flnd. 


Occurrence, habitat. Nymphs of this genus were found in a 
greater variety of situations than any other group. While 
it is for the most part a genus which belongs to the small rills 
it also takes advantage of the secluded places in the larger 
streams. Two species were common in this locality. These 
were Leptophlebia mollis, common after May 20, and L. prae- 
pedita. collected frequently after May 29. 

1911] May-Flies of Fall Creek, N. Y. 109 

Leptophlebia praepedita Etn. 

This species has already been noted by Prof. Needham (Bull. 
86, N. Y., State Mus.) This species is diurnal. Companies 
of them were seen dancing in bright sunlight on the afternoon 
of May 29, June 3, 20 and 21. A mating flight on May 29 
occurred about five o'clock just above a dashing water-fall of 
Fall Creek. As the swarm rose and fell at alternate heights of 
ten to fifteen feet their silvery wings and bodies shone in the 
sunshine like falling snow-flakes. After half an hour of con- 
tinuous flying and soaring the swarm gradually disappeared. 
Of the specimens captured all were males. Earlier in the after- 
noon a similar but smaller swarm was seen flying above a small 
tributary rill. This swarm did not at any time fly higher than 
six feet above the water. Practically the whole swarm was 
captured at one sweep of a large net. Of the captured insects 
forty were males and one was a female. 

Choroterpes basalis Banks. 

This species is plentifully represented in the Creek. The 
nymph is described in Bull. 86 of the N. Y. State Museum. 
Little attention was given to this species beyond the collection 
of nymphs which were constantly associated with Heptagenia. 


This genus is nearly as diversified in habit as Leptophlebia. 
The nymphs have been found in pipe drains, in the gravel and 
trash of still pools, beneath leaves in springs, in the border 
waters of the creek and in its swiftest ripples. Like Lepto- 
phlebia they have been found in rather small numbers and 
widely scattered. My dates for rearings and captures range 
from the second week in May to the last of July. Better 
results would have been obtained if attention had been paid to 
this group earlier in the season for some species were very rarely 
found by May 10th. 

Ephemerella serrata sp. nov. 
Occurrence, habitat. The small nymph of this species was 
found occasionally in restricted areas of the upper Creek where 
it crawled about on stones, or in the trash, which was washed 
by running water. It was found in similar situations at Shef- 
field, Mass. My reared specimens of this species are dated 

110 Annals Entomological Society of America [Vol. IV, 

June 3rd. Collections of the nymphs were made June 12, 14, 
and July 1. By July the nymphs were becoming scarce, 
several cast skins were found on the dry stones of the shore, 
after the last date, but no nymphs were seen. 


Length of body Length of setae 

Male 5 m. m. 6 m. m. 

Female imago 4. .5 m. m. 

Nymph 5 m. m. 1.2 m. m. 

Male imago. General color brown, paler on the legs and below. 
Head, above, eyes very prominent, upper division reddish brown, lower 
one darker. Front of head light, antennae light brownish; ocelli white, 
the lateral ones with an inner lunate band of brown, the median one 
with a complete ring of brown. Thorax, above yellowish shaded with 
brown; mesonotum with its posterior margin edged with brown. Ridges 
of the pleurae shaded with brown. Stemae pale with sub-quadrangular 
areas of brown before the middle legs and a median shield of brown be- 
hind them. Legs pale with a transverse band of brown at the distal 
end of the femora; first tarsus of the foreleg but slightly longer than the 
second; the third twice as long as the fourth. Wings hyaline, brownish 
at the base. Abdomen, brownish above, pale below. Setae 3, pale, 
the joinings of the basal segments ringed with brown. 

Female imago. Eyes small and distant showing the light colored 
occiput between. Body more robust and abdomen slightly darker than 
that of the male. Foreleg twice as long as that of the male. 

Nymph. Small with head and thorax rounded and the setae curv- 
ing fon\'ard o\'er the abdomen. General color a muddy yellow with 
darker markings on the dorsal side, pale below. This species is easily 
distinguished from Ephemcrella deficiens by its double median row of 
spines on tergites 4-7 and by its generally lighter color. Head smooth, 
(PL VI, fig. 5) ; antennae light brownish, first and second basal segments 
edged with brown. Thorax, prothorax with a tubercle like elevation 
on either side the median line. Legs (PI. VII, fig. 2) with the femora 
rather stout and their hinder margins bordered with a row of stout hairs 
sparsely distributed. Claws serrate (PL VII, fig. 2) with a chitinous 
ridged plate on the underside of the tarsus (PL VII, fig. 2). Abdomen, 
above, segments 5-6 pale marked with brown pencilings, other segments 
brown with darker edges. A double row of irregidarly triply dentate spines 
extending over segments 4, .5, 6 and 7. The lateral margins of segments 
4-9 spinose with their posterior lateral angles becoming more acuminate 
posteriorly. Gills present on segments 3-7; Eh-troid lamella absent; 
the superior lamella simple; the inferior fimbriate lamellifonn. Abdo- 
men, beneath, pale with a median row of distinct linear brown spots on 
stemites 1-9. Setae, 3, sparsely beset with coarse hairs; color, pale with 
a transverse band of brown across the center. 

1911] May-Flies of Fall Creek, N. Y. Ill 

Ephemerella deficiens, sp. nov. 
Occurrence, habitat. Of similar habitat and closely asso- 
ciated with Ephemerella serrata in rapid waters, this species 
is the more common of the two and was collected frequently 
during the first half of May. It closely resembles E. serrata 
in shape and size, but the whole body is blackish while the gill 
lamellae, the legs, and the setae are nearly white. 


Length of body Length of setae • 

Male imago 5 m. m. Setae lost 

Nymph 5.2 m. m. 2.-5 m. m. 

Male imago. General color blackish, thorax blackish brown. Head, 
eyes prominent, upper division reddish brown, lower division blackish 
brown (living specimen); ocelli white; antennae, carina and rings 
of ocelli brown. Thorax, above, pronotum brown; the mesothorax 
and metathorax blackish brown with blackish edges; pleurae brown with 
edges blackish; sternum brown with a broad transverse band of yellow 
behind the first pair of legs. Legs nearly white; coxae and an indefinite 
band at distal end of the femur brown. Wings hyaline shaded with 
brown at the base. Abdominal segments shaded with brown giving the 
effect of annular bands dark on the dorsum, paler beneath. Setae 3, 
pale at the base. (Only the bases of the setae remained when the 
specimen was taken from the cage). Forceps and penes are figured in 
PI. IV, fig. 4. 

Nymph (PI. VI, fig. 4, PI. VII, fig. 4, PL VIII, fig. 4). Color black- 
ish brown, femora brown, gills, tibiae and tarsi whitish. Bod}' broadest 
at the metathorax, the thorax arched, the abdomen slightly depressed 
and curving upward at the posterior end. Setae held upright or curved 
over the abdomen. Head, bluntly wedged shaped; ocelli inconspicuous 
whitish; antennae pale with a band of brown just above the basal seg- 
ment. The maxillae with their palpi totally absent (PI. VIII, fig. 4). 
Thorax, above blackish brown, the prothorax bordered laterally by a 
pale longitudinal band with a brown spot at its center; the prothorax 
and mesothorax with a longitudinal stripe of pale yellowish on either 
side the median line. Legs (PI. VII, fig. 4) with the coxae and femora 
brown, the tmder side of the latter with a distinct hook shaped area of 
whitish. Tibia and tarsi pale whitish, the first two pairs of tibiae with 
a broad middle and a narrow proximal band of brown; the tarsi of all 
the legs with a middle band of brown and with the claws similar (PI. 
VII, fig. 4). Abdomen above dark blacldsh brown.' The posterior 
lateral angles of segments 9-10 pale whitish; the lateral margins of seg- 
ments 1-9 spinose serrate with their posterior lateral angles produced 
into fiat spines which become more acuminate posteriorly; segment 10 
without spines and with its posterior margin truncated. Gills present 
on segments 3-7; Elytroid cover absent; gill lamellae whitish shaded 
with brown at the base, superior lamina entire, the inferior one bifed. 

112 Annals Entomological Society of America [Vol. IV, 

Abdomen, beneath, brown without markings. Setae 3, with a circlet 
of spines at their joinings. A broad band of brown across the two outer 

Ephemerella lata, sp. nov. 
Occurrence, habitat. This species was first taken in Sandy 
River, Me., a stream similar to Fall Creek, where it was very 
common. Attempts to rear these nymphs were unsuccessful 
in both Sandy River and Fall Creek therefore a description of 
the nymph only can be here included. It occurs in the most 
rapid water of the stream crawling on the stones much like 
Heptagenia. My dates for its capture in Fall Creek are 
June 20 and 24. It is apparently a rare species in the stream. 

Measurement.— Nymph. Length of body 7.2 m.m. Length of 
setae 3.5 m.m. 

Nymph (PI. VI, fig. 6 ; PI. VII, fig. 1 ) . General color brown with the 
prothorax and the eighth tergite conspicuously white. The fore femora 
very stout (PI. VI, fig. 6) and edged with stout spines. Body arched 
above. The ventral side of the body and the legs flattened and modified 
for clinging to smooth surfaces. Head, sub-quadrangular with the 
broad truncate ledge projecting forward above the rounded heavily 
fringed frontal border (PL VI, fig. G). Antennae with a conspicuous 
triangular ledge projecting above their bases. (Mouth-parts are figured 
on PL VIII, fig. 1). Thorax, prothorax two-thirds as long as the thorax, 
conspicuously whitish but with the posterior ]3ortion shaded brownish, 
the degree varying in difi'erent ages and individuals; mesothorax with- 
out markings; metathorax concealed from above. vStemum flattened, 
brown with pale suture lines. Legs (PL VII, fig. 1) with the femora 
pale marked by a jjale transverse band; tibiae brown with two pale 
transverse bands. Abdomen, rounded above, flattened below. Gills 
on segments 3-S. Elytroid cover absent. Segments 4-9 with their pos- 
terior lateral angles produced back^vard into flat spines; segments 5-S 
with their lateral margins spinose serrate; segments 4-7 with a double 
median row of small tubercles which arise near the posterior border. 
Stemite 9 produced backward into a median rounded lobe and two lateral 
flat spines. Setae 3, light brownish ringed at their basal joinings with 
darker brown. 

Ephemerella tuberculata, sp. nov. 

Occurrence, habitat. But a single specimen of this species 
has been taken. This was a nymph captured on June 22 in the 
gently flowing border water of the upper Creek. 

Nymph, measurements. Length of body, 10.5 m. m.; length of 
setae, 4.5 m. m. 

Body stout arched above, flattened below. Color above dark, 
below pale, with a median double row of distinct brown spots. Fore- 
femora very wide (PL VII, fig. 5). The head (PL VI, fig. 2) with two 

1911] May- Flies of Fall Creek, N. Y. 113 

large erect tubercles on the occiput. Head, sub-quadrangular slightly 
flattened with the head projecting forward; a triangular ridge with the 
median ocellus at its apex projecting above the slightly indented fringed 
frontal margin. (PI. VI, fig. 2). Left maxilla figured on PI. VIII, 
fig. 3. Thorax, more than half the length of the trunk ; color dark above, 
ail except the prothorax pale below; prothorax, above slightlj' arched and 
flaring at its postero-lateral angles, a tubercle at the middle of the lateral 
margin and a smaller one on either side the middle of the posterior mar- 
gin; mcsothorax with one median tubercle. Legs, with femora unevenly 
brown above, pale below; tibiae brown with a pale transverse band 
through the middle; tarsi brown with a pale transverse band at the 
proximal end. Fore-femur (PI. VII, fig. 5) shorter and thicker than the 
others and with its anterior margin unevenly toothed. Anterior edges 
of the other femora entire. Upper surfaces of all the femora with wart- 
like elevations; posterior edge of the first femur and anterior and pos- 
terior edge of the other femora with a row of sparse hairs. Abdomen 
with gills present on segments 3-S, without Elytroid cover, superior 
lamina entire; inferior lamina bifid fimbriate. Segments 2-7 with a 
median double row of spines; posterior margins of segments 1-7 and 
9-10 edged with short hairs; posterior margin of segment 8 with numer- 
ous longer hairs. The posterior lateral angles of segments 3-10 pro- 
duced backward into flat-pointed spines. Setae 3, with numerous hairs 
on their outer and inner margins. 

Ephemerella rotunda, sp. nov. 
(PI. VI, fig. 1; PI. VII, fig. 3; PI. VIII, fig. 5; PI. IX, fig. 6.) 
Occurrence, habitat. This species was taken in portion of 

Pleasant Brook, where there was little water and that strongly 

tainted by pipe drains. But four nymphs were captured. 

The two which were successfully reared proved to be females. 

The dates for their rearing were June 8 and 10. 

Length of body Length of setae 

Female imago 10.5 m. m. 14 m. m. 

Female subimago 10 m. m. 10 m. m. 

Njntnph 10.2 m. m. 6 m. m. 

Female imago. Thorax luteus; legs luteus or whitish; abdomen 
brown; setae luteus with very distinct brown joinings. Head parch- 
ment color. Thorax, above, luteus slightly darker on the mcsothorax; 
pleurae luteus to whitish with brown edges. Axillary cords (Snod- 
grass, '09, The Thorax of Insects and the Articulations of the Wings, 
p. 553) of the fore-wing prolonged into slender acute spines which pro- 
ject baclavard on either side of the hinder lobe of the mcsothorax. 
Axillary cords of the hind wings prolonged in similar but less prominent 
spines. Wings hyaline, costal region sub-hyaline (PI. IV, fig. 6); 
abdomen brown, pale at joinings and beneath. Color evidently mostly 
due to contained ova. Stemite one longer than those following; ster- 

114 Annals Entomological Society of America [Vol. IV, 

nite 7 with its posterior portion overlapping stemite S and with its pos- 
terior margin bilobed. Setae 3, nearly equal length, pilose; color 
pale luteus with distinct brown rings at the joinings. 

Female subimago. Wings sub-hyaline and veins brown. Seg- 
ment 7 not bilobed as in the imago. 

Nymph (PL VI, fig. 1; PL VII, fig. 3; PL VIII, fig. 5). Head and 
body rounded and smooth, and without tubercles. Legs small in com- 
parison with the size of the body. Head, rounded and without eleva- 
tions (PL VI, fig. 1). The left maxilla is shown on PL III, fig. 5. 
Thorax, above smooth. Color, mottled brownish without definite 
markings. Prothorax wider than the head its lateral margins slightly 
flaring, and its width equal to that of the mesothorax; metathorax con- 
cealed from above. Legs weak, the femora poorly developed and with 
a row of hairs on its posterior margin (PL VII, fig. 3). Abdomen, above 
rounded, gills present on segments 3-7, the posterior margins of segments 
4-9 with a double median row of small spinose elevations; segments 3-9 
with their posterior lateral angles produced into flat spines whose mar- 
gins are spinose serrate. Setae brownish indefinitely banded with 
whitish; basal joinings with circlets of hairs. 

Ephemerella cornuta, sp. nov. 

Occurrence, habitat. This, before mentioned species, was 
reared at Salisbury, Connecticut. The dates given by Miss 
Smith for its capture and rearing are July 20, 21. But two 
stages, those of the male sub-imago and the nymph are repre- 

Length of body Length of setae. 

Male subimago 10.5 m. n. 

Nymph 10 m. m. 6. m. m. 

Male subimago, general color pale luteus with annular bands of 
brown on the abdomen. Eyes prominent. Head with ocelli white, 
encircled with broad bands of blackish brown; carina, frons and occiput 
pale; antennae light brown. Thorax, above pale luteus; the prothorax 
irregularly streaked with brown. Axillary cords of the fore-wing 
produced into a slender point extending backward on either side the 
median lobe of the metathorax, such prolongations not evident in the 
hinder wing. Legs, pale, the fore femora lightly shaded with brown. 
Wings, sub-hyaline, brownish at the base (PL IX, fig. 5). Abdomen 
pale whitish, with annular bands of brown shading. Setae 3, whitish. 

Nymph. Body slender, and tapering from the mesothorax. Fore 
femora shorter and wider than the others and with its anterior margin 
unevenly toothed (PL VII, fig. G). Head, with prominent incurving 
horns just below the antennae; general shape sub-quadrangular with 
the posterior angles rounded, and the frontal margin fringed with hairs 
and projected forward; origin of the antennae partly hidden by the 
curving ledges at the bases of the horns (PL VI, fig. 3). (Left maxilla 

1911] May-Flies of Fall Creek, N. Y. 115 

shown on PI. VIII, fig. 6.) Thorax, mottled brownish; prothorax 
quadrangular, its angles closely fitting to the mesothorax. Legs with 
the margins of the second and third femora entire; the first femur fig- 
ured on PI. VII, fig. 6. Abdomen, rounded above, flattened below; 
without dorsal spines or tubercles. Gills on segments 3-7; postero- 
lateral angles of segments 3-9 produced into flattened spines becoming 
more acuminate posteriorly; posterior margins of segments 2-10 sparsely 
edged with hairs; lateral margins of segments 3-9 spinose serrate. 
Setae, pale, except for a single brown ring at the base of each. 

Tricorythus allectus Needham. 

This species was described by Prof. Needham in Bull. 86, 
N. Y. State Mus. as Caenis allecta, but afterward referred by 
him to the genus Tricorythus, in Bull. 124 N. Y. State Mus. 

Occurrence, habitat. These nymphs are closely associated 
with Caenis, though they have not thus far been often found in 
the ill smelling mud generally preferred by that nymph. They 
clamber about in fine silt and sand, and the particles which 
adhere to their hairy bodies make them practically invisible. 
A handful of mud which appears to contain no sign of life, will 
after a few minutes draining, reveal slowly moving bits of mud 
which may prove to be either Caenis or Tricorythus according 
to the quality of the mud. Tricorythus allectus is one of the 
commonest species in Fall Creek. During June and July, the 
imagoes may be found strewn upon the surface of little protected 
inlets along its shores, or caught in the meshes of the spider- 
webs on walls and bridges near it. 

Measurement. Length of body 6.5 m. m. Length of setae 4 m. m. 
Mouth — parts and gill lamella (PL XI.) The nymphs of 
Tricorythus may be at once distinguished from those of Caenis 
by the shape of the elytroid gill cover, rounded at the 
end in Caenis, distinctly triangular in Tricorythus (PL XI, fig. 1). 
Color yellowish, pale below; abdomen marked with transverse bands 
of brown broken by a median longitudinal pale stripe. Elytroid 
lamella prominent, purplish brown at the base. Body all over sparsely 
beset with hairs. Antennae pale with basal segments brown. Legs 
pale with a blackish spot at the proximal joint of each tibia. Gills not 
wholly hidden by elytroid lamellae. Lateral spines on segments 2-9. 
Setae 3. 


This genus is plentifully represented by Caenis hilaris, Say, 
and by nymphs of some other species not yet reared here. 

116 Annals Entomological Society of America [Vol. IV, 

Chirotonetes albomanicatus Needham. 
Occurrence, habitat. Nymphs of this abundant species 
were found full grown in the dashing waters of the falls and riffles 
from the latter part of May to August 12th. In May they were 
seen to emerge in greatest numbers during the late afternoon 
and twilight. They crawled up on the shore leaving their cast 
skins clinging to the stones or less often they flew up directly 
from the mid current. The dark wings and body and the white 
forelegs of the sub-imago made it very conspicuous as it rested 
upon the gray stones or flew upward. Robins made a regular 
custom of coming to the shores and collecting the insects as they 
emerged. From four to six nymphs might be often seen pro- 
jecting from their beaks. 

Siphlurus alternatus Say. 
This elegant species has been found to be very desir- 
able for the indoor aquarium. It lives in still pools and 
demands only a minimum supply of fresh water and plenty of 
plant food. It was common all through April, May, and a 
portion of June. Nymphs kept in the laboratory aquarium 
nearly all lived to emerge and doubtless could have been kept 
there several months had they been taken early enough. 

Baetis pygmea Hagen. 

This species has been found chiefly interesting for its habits 
of egg-laying. From early June to late August the stones in the 
waters of the creek were covered with small elongate egg- 
patches rounded at one end, narrower and sharply' squared off 
at the other. Plate XII, fig. 1 shows a stone about seven by 
ten inches in size on which the egg masses were scattered with 
average abundance. The surface upon which the eggs rested 
was the down-stream side of the stone and that portion where 
they were thickest was nearest to the surface of the water. On 
the same plate (fig. 2) is shown a photograph in which the 
patches are enlarged sufficiently to show the individual eggs. 
The laying of the eggs may be seen if one closely watches some 
stone which is marked as a favorite site by the presence of many 
masses. The following observation was made through an ordi- 
nary reading glass, but the processes may be easily seen with 
the naked eye. 

Flying close to the surface of the water, the insect alighted 
on a stone projecting slightly from the water and well protected 
from the force of the current on its downstream side. She 

1911] May-Flics of Fall Creek, N. Y. 117 

immediately walked to the protected side and downward to the 
water. First, wrapping her wings about the abdomen, she 
made several attempts to immerse her head and thorax. This 
appears to be the critical stage of the performance, for many- 
females are washed from the stone while attempting it. Once 
beneath the surface she started on a tour of inspection for the 
proper surface. This tour lasted for several minutes during 
which time she continually walked to and fro, pausing, feeling 
with the abdomen, and passing on unsatisfied. When a suit- 
able place was finally found she braced her legs firmly, bent the 
abdomen downward, curved the setae upward and pressed 
the openings of the oviducts closely to the surface. The whole 
abdomen was then swung from side to side with a slow pendu- 
lum-like motion, each stroke leaving an irregular row of minute 
white eggs adhering to the surface. The strokes were at first 
■somewhat circular and longer than those which followed. As the 
egg mass grew in length the insect moved forward a little to allow 
the eggs to lie in succeeding rows. When the egg supply was ex- 
hausted she stopped with a jerk of the abdomen and proceeded 
to clamber out of the water. When examined in the laboratory 
the abdomen of this female was found to contain only a few 
undeveloped eggs in the ovaries and none in the oviducts. 

Ameletus ludens Needham. 
Occurrence, habitat. In Pleasant Brook on April 25 
nymphs of Ameletus ludens, were found in great abundance 
resting upon the bottom or darting about much like the nymphs 
■of Callibaetis. They were the dominant insect of the stream 
at this time. One soft bottomed pool about four feet long, 
and two wide yielded about 300 nymphs in half an hour's 
collecting, and many more remained. Full grown nymphs 
were placed in cages for purposes of rearing. The female sub- 
imago was found to correspond with Prof. Needham's descrip- 
tion given in Bull. 86 of the N. Y. State Mus. Attempts were 
then made to secure a male to add to the life-history since 
Prof. Needham was unable to procure one. The results of the 
rearing were as follows: 

April 29, 1 nymph emerged, transformed April 30. Female. 
April 30, 1 nymph " " May 1. " 

May 7, 1 nymph " " May 8. " 

May 8, 3 emerged nymphs, " April 9. " 

Repeated rearings failed to secure a male specimen. An 
•enclosure was then made in a neighboring rill by means of boards 

118 Annals Entomological Society of America [Vol. IV, 

and fine meshed wire-cloth. Over this, a cheese-cloth tent was 
erected, and in it a large number of nymphs were placed. The 
records of the results obtained are as follows : 

May 8, 12 nymphs emerged before 12 M., transformed before May 9, 12 M. 
All Females. 

May 10, 40 " " before 12 M., " before 12 M., May 9. 

All Females. 

May 12, 25 " " before 12 M., " May 13, before 12 M. 

All Females. 

That these nymphs emerged so regularly before noon was 
doubtless due to the fact that the sunshine reached the tent only 
at this time and in the late afternoon. Rearings were made as 
long as the season lasted but neither among the reared specimens 
nor among the nymphs collected could a male be found. A few 
specimens of the same species were also collected in two other 
streams near Ithaca, but no males were found. The fact that 
no male specimens have been taken neither in collecting, or 
rearing, indicates a case of parthenogenesis in this species. 

Female imago. Measurements. Length of body 10 m. m. Length 
of setae 10 m. m. Color bright reddish brown with whitish areas on 
the thoracic pleurae. Abdomen reddish brown slightly paler beneath 
with the ventral ganglia marked by darker areas. Antennae brown, 
paler at the base, the second segment very long. Thorax brown; legs 
brown, the third pair slightly paler than the others; wings hyaline with 
their bases shaded with brown and the veins very distinct. Setae brown 
with their joinings distinctly lined with brown. 


1. The physical features of Fall Creek make possible the 
greatest variety of aquatic conditions. A study of the abundant 
May-fly fauna which lives under these conditions has revealed 
a series of striking adaptations to environment. 

2. One may best observe the nymphs and secure adults of 
uncommon species by rearing the insects in their own sur- 
roundings. For this purpose a new type of breeding cage has 
been described and figured. 

3. Observations upon representatives of 17 different genera 
have shown some interesting points in structure and ecology. 
Among these have been described the structural adaptations 
of various nymphs, the striking peculiarities of Ephemerella 
nymphs, the emerging of Blasturus cupidus, the swarming of 
Ephemera and Leptophlebia, and the egg-laying of Baetis. 

4. No male specimen of Ameletus was secured either by 
collection or among 83 reared specimens. I have, therefore, 
suggested that this species may present a case of parthenogenesis. 

1911] May-Flies of Fall Creek, N. Y. 119 

EXPLANATION OF PLATES. (All figures much enlarged.) 
Heads of Ephemerella nymphs. 
Fig. 1. Ephemerella rotunda sp. nov. 
" 2. " tuberculata " 

" .3. " comuta " 

" 4. " deficiens 

" 5. " serrata " 

« 6. " lata 

Legs of Ephemerella Nymphs. 
Fig. I. Right legs of Ephemerella lata sp. nov. 

" 2. " foreleg " " serrata " 

" 3. " " " " rotunda 

" 4. " " " " deficiens 

" 5. " " " " tuberculata " 

" 6. " " " " comuta 

These drawings are all made from the upper or dorsal aspect. The enlarged 
sketches of the claw and tansus of each show the ventral aspect of the tarsus with 
its chitinous comb which is used in clinging to the rocks. 

Mouth-parts of Ephemerella nymphs. 
Fig. 1. Mouth-parts of Ephemerella lata. sp. nov.; r. md., right mandible; 
1. md., left mandible; Ir., labrum; 1., labium; mx., left maxilla; hy., hypopharynx, 
viewed from above. 

Fig. 2. Left maxilla of Ephemerella serrata sp. nov. 
" 3. " " " " tuberculata " 

" 4. " " " " deficiens 

"5. " " " " rotunda 

" 6. " " " " comuta " 

Epeorus and Ephemerella. 

Fig. 1. Epeorus humeralis sp. nov. Portions of the 3rd and 4th abdominal 
segments of the nymph with gills removed, to show lateral spines. 

Fig. 2. Forceps and penes of Epeorus humeralis, sp. nov. (From below.) 
" 3. " " " " Ephemerella serrata sp. nov. (From below) 

« 4. " " " " " deficiens " (From below). 

" 5. Wings of " comuta " 

" ^ " " rotunda 

" 7. " Epeorus humeralis " 

Iron fragilis sp. nov. 
Fig. 1. Mouth-parts of the nymph; 1, outer aspect of the labium; IL, inner 
aspect of the labium; r. md., right mandible; 1. md., left mandible; hy., hvpopharynx. 
Fig. 2. Wings. 

Fig. 3. Foreleg of the male imago. 

Fig. 4. Forceps and penes from below; dotted lines represent the portion of 
the penes hidden by the last stemite. 

Tricorythus allectus Needham. 
Fig. la. Elytroid lamella; hy., hypopharynx; 

" Ir. labrum; r. md., right mandible; 1. md., left mandible. 
" mx. left maxilla; 1., labium. 

Fig. 1. Surface of stone covered with masses of Baetis eggs. 
" 2. A few of the masses enlarged. 

Annals E. S. A. 

Vol. IV, Plate VI. 

G.H hr.rj'" 

Annals E. S. A. 

Vol. IV, Plate XVII. 

O.-H h,„j,„ 

Annals E. S. A, 

a H hl^rj"^ 

Annals E. S. a. 

Vol. IV. Plate IX. 

G. M/f!„j,^ 

Annals E. S. A. 

Vol. IV, Plate X. 

Ch fv,.}'^ 

Annals E. S. a. 

Vol. IV, Plate XI. 

Ci,H- ^erjun. 

Annals E. S. A. 

Vol.. IV. Plate XII. 

A. H. Morgan. 


By Charles H. T. Townsend, 
Piura, Peru. 

The work on the female reproductive system, eggs, and 
first-stage maggots of the Tachinid fiies and their alUes, begun 
in 1908 by the writer at the Gipsy Moth Parasite Laboratory 
in Massachusetts, under the direction of Dr. L. O. Howard, 
Chief of the Bureau of Entomology, has been prosecuted to 
date as time permitted. The results are now such that it 
becomes desirable to make an announcement of them in brief. 
This announcement is in advance of a series of much more 
complete papers, which will contain plates of the female repro- 
ductive and accessory organs, eggs, first-stage maggots, and 
cephalopharyngeal skeletons of the latter, some 200 drawings 
having already been completed for this purpose. 

The female reproductive and accessory organs in the Mus- 
coid flies consist of (1) ovaries, (2) oviducts and common ovi- 
duct, (3) spermathecae and their ducts — 3 in number, (4) tub- 
ular (or coUeterial or accessory) glands and their ducts — 2 in 
number, (5) uterus when present, including what may be 
termed the preuterus which is present in some forms, (6) uter- 
ovagina, being a vagina proper which functions anteriorly 
as a true uterus when latter is absent, and (7) ovipositor or 
larvipositor and appendages. 

The functions of most of the above organs are generally 
understood, but the following points need mention: 

The tubular glands function as secretory organs for the pro- 
duction of the viscid fluid for coating the eggs, and are more 
or less rudimentary in those forms that deposit maggots; the 
preuterus is a small sac at the head of the uterus, in which the 
egg of some forms is fertilized before passing into the uterus 
proper, the spermathecal ducts opening into it ; the uterovagina 
is a short tube homologous with the so-called insect vagina, 
its anterior portion filling the office of uterus in those forms 
without distinct uterus, the spermathecal and tubular gland 
ducts opening therein, its posterior end filling the office of 
vagina. The openings of the spermathecal ducts always mark 
the transition from common oviduct to functional uterus. 


128 Annals Entomological Society of America [Vol. IV, 

The results so far secured in the present work indicate at 
least 37 distinct series in the Muscoid flies (exclusive of Antho- 
myiidae and Acalyptratae), based mainly on the characters 
of the reproductive and accessory organs of the female fly, 
the egg, first-stage maggot, and in some cases the facial plate 
and other characters of the external anatomy of. the adult. 
Undoubtedly further work will demonstrate the existence of 
further series demanding recognition. Briefly the series so 
far recognized may be tabulated as follows: 

1. Trichopodine series — Xanthomelanodes peruanus n. sp. 
(Peru) dissected and drawn, TD 3983. No uterus, utcro- 
vagina short and broad, tubular glands short and thick, sper- 
mathecal ducts very long and spermathecae attached in hood, 
oviducts of moderate length and thickness, ten eggtubes in 
each ovary in the above species, eggs flattened and pink-sal- 
mon to flesh-brown in color when mature. The chorion of egg 
is beautifully honey-comb reticulate. Xanthomelanodes and 
allies. Trichopoda and allies, many of which have been 
dissected, have same eggs and ovaries, and almost certainly the 
same type of reproductive system. 

2. RuTiLiiNE series — Rutiha sp. and Amphibolia sp. (Aus- 
tralia) dissected, TD 18G4, 1866. Uterus present, but its 
character not yet known. Maggots long and slender, hairy 
or furnished with hairs at anal end. Rutilia maggot has anal 
hairs, Amphibolia maggot is thinly hairy on body. Rutilia, 
Amphibolia and allies — Australian flies, most of rather large 
size. These will probably need division into several series. 

3. Phasiine series — Uterus present, form not known. 
Egg very elongate, slender, TD 480 (South Carolina), near 
Alophora, has what seems a piercing larvipositor or ovipositor, 
but curved in the opposite direction from that of Compsilura 
and not so sharp apically. Phasia, Alophora, Hyalomyia and 
allies, but these have yet to be studied. 

4. Graphogasterine series — Hyalomyodes sp. (South 
Carohna) dissected, TD 481. Uterus present, eggs and 
maggots slender. Hyalomyodes and alhes, and probably 

5. Glossinine series — Functional uterus, whether uterus 
proper or uterovagina, greatly enlarged to hold the maggot 
until fully grown and ready to pupate, some special provision 
being evidently present for the feeding of the maggot during 

1911] . Study of Muscoid Flies 129 

its three stages. A most remarkable and distinct type, in any 
event, not only in its reproductive system and habit, but also 
in its venation and other characters. Glossina and allies — 
African blood-sucking flies, carriers of Trypanosomae of var- 
ious forms of sleeping sickness in man and animals. 

6. Stomoxydine series — Stomoxys calcitrans (Peru) dis- 
sected and drawn, TD 3985. No uterus, uterovagina short, 
spermathecal ducts long and doubled, tubular glands fairly 
well developed but not longer than oviduct plus common ovi- 
duct. Eggs elongate and deposited on dung. Adult with 
piercing mouthparts in both sexes. Ovipositor with a dorsal 
pair of bristly, slightly curved, subcylindrical chitinous processes. 
Stomoxys, and j^robably Lyperosia, Haematobia and allies — 
blood-sucking flies and probable carriers of microzoa of certain 
cattle diseases. 

7. Calliphorine series — Comp-somyia macellaria (Peru) 
dissected and drawn, TD 3984. No uterus, uterovagina very 
short; spermathecal ducts very short, only as long as the sper- 
mathecae themselves; tubular glands only very moderately 
developed, about as long as common oviduct plus oviduct, the 
latter hardly half the length of the former. Ovipositor simple. 
Mouthparts fleshy. Eggs elongate, deposited in sores or on 
meat, the product of both ovaries being deposited at one time. 
Compsomyia and allies, and probably Calliphora, Lucilia and 

8. Mesembrinine series — Probably a distinct series comes 
here, including Mesembrina and allies, and especially Dr. 
Adolf Lutz's strange Brazilian fly which Prof. Hermann deter- 
mines to be Pseudogametes, and which seems to have Mesem- 
brinine affinities. 

9. Muscine series — Musca domestica (Peru) dissected 
and drawn, TD 3982. No uterus, uterovagina elongate; a 
pair of uterovaginal pouches springing from lateral anterior 
walls, one on each side below insertion of spermathecal and 
tubular gland ducts, being accessory copulatory vesicles of 
Hewitt; tubular glands long and slender; eggs elongate, depos- 
ited on dung, etc. Musca and allies. 

10. Sarcophagine series — Sarcophaga 2 spp. (Peru) dis- 
sected and drawn. Uterus, when distended, heart-shaped or 
cordate, maggots rather irregularly disposed therein. Utero- 
vagina short, but with two dorsally-lying sacs or large pouches 

130 Annals Entomological Society of America [Vol. IV, 

developed from its anterior ventrolateral walls, one on each 
side, these two sacs and the uterovagina together forming the 
heart-shaped functional uterus, which contains the eggs until 
the embryo has developed to the fully formed maggot ready for 
deposition on host or food-substance. Sarcophaga and allies; 
and possibly Rhinophora, Melanophora, Brachycoma and 
allies, the last three genera being included by external anatomi- 
cal analogy. Brachycoma is a Bombus inquiline or parasite; 
Rhinophora and Melanophora are terrestrial-isopod parasites. 

11. Metopiine series — Metopia sp., TD 3988; Selenomyia 
sp., TD 3998; Sarcomacronychia sp., TD 3996 (all Peru) 
dissected and the first drawn. Uterus thick short V-shape, 
the arms of the V being productions anteriorly of the ventral 
walls of the uterovagina on each side, on the same plan as that 
of Sarcophaga but in different form, not rounded but rectangu- 
lar pouches, the form doubtless due largely to regular disposi- 
tion of the contained eggs and maggots; spermathecal ducts 
very long, doubled on themselves; tubular glands thick and mod- 
erately long. Maggots and eggs regularly arranged on end in 
uterus in triple and quadruple file. The uterovagina forms the 
base of the V, thus functioning as part of the uterus; the empty 
arms appear as blind tubes. Metopia, Selenomyia, Sarcoma- 
cronychia and allies. It is to be noted that the facial plate in 
the adult of Metopia is widely different from that of the rest 
of the group, probably due to antennal development, thus indi- 
cating its inferior rank in this series. Largely muddauber- 
wasp inquilines or parasites, feeding on contents of nests. 
Related to the Sarcophagine series. 

12. CoMPSiLURiNE series — Compsilura concinnata (Eu- 
rope), Dexodes nigripes (Europe), Vibrissina sp. (Florida), 
and Eucelatoria spp. (Florida and Peru) dissected; TD 290, 
132, 775, 1229, 3906. Uterus slender, long, in several coils, 
maggots and eggs normally obliquely on end, the maggots 
usually in single file, deposited subcutaneously in host through 
the hollow curved piercer of female fly, the single file arrange- 
ment doubtless for the purpose of facilitating the passage of 
maggots through the piercer. This type is a modification of the 
Hemimasiceratine type, with the addition of a piercing larvi- 
positor for subcutaneous deposition of the maggots. Comp- 
silura, Vibrissina, Eucelatoria and allies, including so far as 
known all flies whose females are provided with a curved 
piercing sharp-pointed larvipositor. 

1911] Study of Muscoid Flies 131 

13. Tachinine series — Tricholyga sp. (Peru) dissected and 
drawn, TD 3971. Tachina spp. (Europe, America and Ja- 
pan) dissected. No uterus, uterovagina normal and capable 
of holding but few eggs at a time ; eggs oval and flattened, pro- 
vided with a terminal dorsal hinged lid or cap for exit of maggot, 
normally deposited in a comparatively undeveloped state of 
embryo on host. Tubular glands very long and highly func- 
tional. Chorion hard, opaque, not reticulate. Tachina, Tri- 
cholyga and allies. Everything here points to egg deposition, 
and it is thus difficult to understand Dr. I. C. Nielson's record 
of larviposition for his Tachina larvarum, female flies of which 
appear to be the same as the ordinary form from which we secured 
deposition of great numbers of eggs but never a maggot, at the 
Gipsy Moth Parasite Laboratory. 

14. Meigeniine series — Eumyothyria sp. (Peru) dissected 
and drawn, TD 39S1. Uterus in a single coil, tubular and 
thickened when full of eggs; egg oval and flattened like that 
of Tachina, maggot developing within egg in uterus certainly 
to some extent, but egg evidently deposited on host after a cer- 
tain period of uterine incubation. The fiat eggs exhibit a 
shingled arrangement in the uterus. The long well developed 
tubular glands show that the eggs are intended for deposition 
as such, before the escape of the maggot. Both tubular glands 
and eggs are practically same as in the Tachinine series. Eumy- 
othyria and quite certainly Meigenia and allies. TD 651 
(Florida) with small brown flat eggs, which were certainly 
uterine, and TD 738 (Ocean Beach, So. Florida), with Plagia- 
like venation and small fiat uterine eggs may possibly come here. 

15th series — Apparently what Coquillett determines as 
Sturmia distincta, which seems same as protoparcis Towns, 
and is probably referable to Zygobothria (Florida), TD 619; 
and other spp. (Europe and Florida) dissected. Short coiled 
strap-like uterus full of maggots and eggs on end, after style 
of Hystriciine series (which follows) but in only two or three 
coils and with white maggots which are evidently deposited on 
hosts, being without anal membraneous pad for leaf-attachment. 
Zygobothria and allies, provided Z. bimaculata Htg. of Europe, 
the type of the genus, agrees herein as it seems to in external 
characters. Type specimens of many genera will have to be 
dissected before we will know what name to give this series. 

132 Annals Entomological Society of America [Vol. IV, 

IG. Hystriciine series — Archytas sp. (Peru) dissected and 
drawn, TD 3989; and Melanophrys, Varichaeta, Copecrypta, 
Echinomyia, Jurinia, Paradejeania, Dejeania, Saundersia, and 
many others dissected and their maggots drawn. Very long 
coiled strap-like uterus, in many coils, band-like, wide and thick, 
full of eggs and maggots on end, the thickness of the uterus 
corresponding to the length of the maggots and eggs contained, 
but the uterus lying on edge in a spiral like a watch-spring; 
maggots developing therein and becoming dark colored when 
mature from the blackish dorsal and lateral minute scale-like 
plates, deposited on the foliage of plants in proximity to their 
hosts, being furnished with an anal membranous attachment 
pad for adhering to plant surfaces. Tubular glands short and 
small. The above named genera and their allies, forming a 
very large and predominant series especially in the mountain- 
ous regions of both Americas, and representing the most recent 
phase of Muscoid fly evolution. This series will need to be 
divided into several groups. 

17. Masiceratine series — Blepharipa politana n. sp. 
(Peru) dissected and drawn, TD .3977; others dissected and 
their maggots and eggs drawn. Very long coiled tubular slender 
uterus, in many coils, filled with thousands of microscopic 
eggs which are held till the contained maggots are fully formed 
when they have become black in color and are deposited on 
foliage to be swallowed by hosts in feeding, being placed in 
proximity to leaf-eating insects, probably always lepidopterous 
larvae. Tubular glands moderately thick and developed. 
The chorion of the egg in this series exhibits always a honey- 
comb-like or network system of reticulation, though often also 
showing minute light-colored points appearing as microscopic 
punctures. The chorion of egg is oval in outline viewed from 
above. TD 877 (So. Florida), the adult of which at first sight 
appears much like Cnephalia, has the egg-substance protruded 
beyond the chorion at both ends, giving the eggs a decidedly 
slender and pointed appearance especially when seen in situ 
through the walls of the uterus. The explanation of this pecu- 
liarity has yet to be learned. The series includes Masicera, 
Blepharipa, etc., forming with other leaf-ovipositing flies a large 
group more especially predominant in the lowlands of both Ameri- 
cas as well as other parts of the world, and representing an extreme 
phase of Muscoid fly evolution somewhat less recent than that 
of the Hystriciine series. 

1911] Study of Mtiscoid Flies 133 

ISth series — Species agreeing in external characters with 
Eumasicera but certainly not that genus (Peru), TD 3987; 
dissected; probably Exorista futilis O. S., determined in the 
adult by Mr. W. R. Thompson (Massachusetts), TD 344, 
361. Uterus short, thick, tubular, in only two or three coils, 
filled with microscopic oval eggs of same character as those 
of Masiceratine series except that they show no honeycomb 
or network reticulation but a concentric-ring or concentric- 
arc pattern viewed from above, apparently due to a dispo- 
sition of ridges or wrinkles in the chorion. Eggs deposited 
on foliage, probably for lepidopterous larvae only. The above 
mentioned forms must, I think, belong to the same genus as 
the reticulation of the chorion is of the same character ; but until 
the type species of Exorista, Sturmia, and various other genera 
are dissected, it will be impossible to say what generic name must 
be given them; and still further genera must be dissected before 
we can know what name to give this series, which may include 
Phorocera, or even older genera. TD 437, which seems refer- 
able to Phorocera on external adult characters, probably comes 
in this series; it is European. 

19th series — Ophirosturmia cincta gen. et sp. nov. (Peru) 
dissected and drawn, TD 4012. Uterus tubular and elongate 
in four to six coils, stouter near head, filled with microscopic 
oval brownish-yellow to yelk)w-brown eggs, whose choria 
exhibit a honeycomb reticulation, the eggs deposited on leaves 
in proximity to certain leaf-eating insects. There is no punc- 
tulation to the chorion. Distinguished from the Masiceratine 
series by difference in spermathecal ducts, and by the brown- 
yellow color of the mature eggs, as well as other points. Apical 
cell ending well before wing-tip. 

There are other series yet to be defined among the leaf-ovi- 
positing forms. 

20. Gymnochaetine series — Gymnochaeta sp. (Peru) dis- 
sected, maggot drawn, TD 3973. Maggots and eggs on end in 
two or three rows in very long subtubular coiled uterus ; maggots 
black by reason of the body segments bearing a large dorsal 
and two small lateral colored plates, these made up of minute 
colored scale-like plates of different form from those of the 
Hystriciine series; the maggot with ventral locomotory spine- 
pads and spine-rows composed of very microscopic spines and 
evidently specially fitted for locomotion in the open; no anal 

134 Annals Entomological Society of America [Vol. IV, 

membraneous attachment pad being present. This is a totally 
new type of maggot and indicates a widely diverse habit from 
others so far known. Gymnochaeta and allies. 

21st series — Ophirion mirabile gen. et sp. nov. (Peru) dis- 
sected and drawn, TD 3980. Maggots and eggs rather loosely 
disposed in long slender tubular uterus, the eggs in two rows 
obliquely on end; an elbow-like preuterus, uterus arising 
from inside angle of elbow, common oviduct opening into one 
end of elbow and the spermathecal and tubular gland ducts 
into the other end; oviducts long and slender, tubular glands 
very short; maggots of a peculiar and new type, remarkable 
in that they appear emarginate laterally and are furnished with 
a dark-colored pattern. 

22. PsEUDODEXiiNE series — Ophirodexia pulchra gen. et 
sp. nov. (Peru) dissected and drawn, TD 3999. Uterus 
thick, tubular, in two coils, with eggs and maggots obliquely 
on end; a true preuterus homologous with the forward half or 
more of the uterovagina, the uterus having evidently developed 
from the walls of the vagina proper; spermathecal and tubular 
gland ducts opening into the preuterus, which has capacity of 
just one egg; all these ducts very short; tubular glands only 
about three times as long as preuterus, or as long as common 
oviduct plus oviduct; oviducts very short and thick; common 
oviduct much longer and slender, being about twice length of 
preuterus. Maggots slender. Abdomen of fly elongate, sub- 
cyhndrical; facial plate long, wide, not constricted by vibrissal 
angles, cut off abruptly below; antennae elongate, arista plum- 
ose. These characters apply to the above form, TD 3999. 
Ophirodexia and allies, including probably Pseudodexia, Lep- 
toda, Cordyligaster, and a host of others. Atrophopoda may 
come here, or may be very distinct. 

23. OCYPTERINE series — Probably a distinct series comes 
here for Ocyptera and allies. The genus has been dissected, 
but not with sufficient care. The eggs are elongate, stout, 
subcylindrical ; maggot unknown to me. 

24. Thrvptoceratine series — Probably a series comes 
here for Thryptocera and allies. Eggs slender, elongate. 
Maggot not known to me. 

25. Hemimasiceratine series — Uterus present, doubtless 
coiled, containing elongate white maggots which must be 
deposited on host. Hemimasicera, Sisyropa and alhes. These 

1911} Study of Muscoid Flies 135 

genera dissected and maggots drawn, but the form of uterus 
was not determined. 

26. SiPHOSTURMiNE series — Siphosturmia sp. (Peru) dis- 
sected. TD 4002. Single-coil uterus, sub-tubular, with two 
rows of eggs and maggots; very long ovipositor, sub-chitinous, 
in its unextended position in abdomen measuring nearly one- 
half the length of uterus; ovaries slender, elongate, with few 
eggtubes; oviducts long, common oviduct same length as ovi- 
duct; spermathecae elongate-oval, two joined in hood, the third 
free; spermathecal ducts elongate, about as long as common 
oviduct ; tubular glands moderately elongate and slender, about 
as long as common oviduct plus oviduct. Siphosturmia and 
allies. Florida specimens of the genus have also been dissected. 

27th series — Anisia, Erynnia, Gymnostylia spp., or forms 
allied to these genera so far as can be known by external adult 
characters, including Hypostena barbata of Coqt. in part 
(Peru and Maryland) dissected, TD 4001, 4.33. Plump white 
maggots with well-developed spine rows on ventral surface. 
Uterus slender, tubular, coiled three times within itself; mag- 
gots in single file, oblique to nearly longitudinal in position 
according to degree of crowding in uterus; spermathecal ducts 
long, tubular glands not long; ovaries elongate, with few egg- 
tubes. The maggots of the Peruvian form (TD 4001) show 
five highly developed ventral spine rows; those of the D. C. 
form (TD 433, collected by Knab on Plummers Island follow- 
ing a beetle of Calligrapha bigsbyana) are very similar, having 
five transverse rows of strong hooked spines on posterior half 
of median ventral surfaces. Maggots of TD 352 (Mass.) 
are short and plump, with strongly marked complete rows of 
spines encircling the body, and may not belong here; the fly has 
a different facies. The series embraces flies that are parasites 
of both the grubs and adults of certain beetles, especially 
Chrysomelidae. The European tachinid parasite of the import- 
ed elm-leaf beetle apparently belongs here. Many genera will 
likewise have to be dissected before this series can be named. 

28. Parexoristine series — Parexorista cheloniae (Europe) 
and Parexorista sp. (Mass.) dissected, TD 432. Uterus in a 
single coil, containing elongate subcylindrical eggs which are 
furnished with a pedicel, the eggs incubated in uterus sometimes 
to an advanced stage of the embryo and then deposited on host 
attached to skin or hairs of latter by pedicel. Parexorista and 

136 Amials Entomological Society of America [Vol. IV, 

29. Dexine series — Almugmyia arida gen. et sp. nov. 
(Peru) dissected and drawn, TD 3979. Dexia, Mochlosoma, 
Sirostoma, Sardiocera, and allied forms dissected and maggots 
drawn. Uterus thick, tubular, in one coil; eggs and maggots 
in a very oblique and somewhat spiral arrangement; ovaries 
very slender, elongate, consisting of few eggtubes; common ovi- 
duct rather long, oviducts short, spermathecal ducts long and 
doubled, tubular glands small. The maggots of Almugmyia 
and Sirostoma have anal spiracular tubes carried out in long 
slender anal processes which are tipped with bristles; the eggs 
of the first are sharply pointed anally, due to the presence inside 
the chorion of the anal bristles of the developing maggot. 
Probably the genera above mentioned all come in this series 
with many others, but the reproductive-system characters 
above given have not yet been verified for the genera other 
than Almugmyia. Parasites of white-grubs, woodboring-grubs, 
and probably others. 

30. Myiophasiine series — Ennyomma globosa (South Car- 
olina and Florida) dissected, TD 509. The maggot is so 
slender as to be almost filiform. Uterus present, but form not 
•determined. Ennyomma, Myiophasia and allies. Parasites 
of weevil grubs in green fruits. 

31. Phasiopterygine series — Phasiopteryx spp. (Colo- 
rado, Veracruz and Peru) dissected, TD 1791, 1791a, 4005. 
Uterus present, very long and slender, in many irregular knot- 
like coils and turns, the upper part more regularly coiled, receiv- 
ing the eggs while latter are still microscopic, the eggs growing 
to full size in upper part of uterus. Ovaries and oviducts very 
small, the latter not over one-fourth the diameter of a full- 
grown uterine egg. Preuterus present. Spermathecal ducts 
very short, tubular glands short. Maggots with five longitudi- 
nal rows of strongly chitinized segmental plates, those of median 
row wide and covering dorsum of segments, those of inner lat- 
eral row narrowed, those of outer lateral row presenting a ser- 
rate outline below; underside white and soft. The chitinized 
plates of the maggot vary from brownish-yellow to black, over- 
lap when the maggot in contracted, the segments telescoping 
and producing a strongly emarginate outline both from above 
and in profile. These plates are evidently ambulatory in func- 
tion as well as protective against conditions of life in the open 
or subopen. Maggots elongate when not contracted, moderate- 

1911] Study of Miiscoid Flies 137 

ly wide, flat below; wide and short when contracted. The habit 
is larviposition, but one can not even guess at the larval habit 
and host relation. This is a most interesting and remarkably dis- 
tinct type. Although the Colorado and Veracruz specimens of 
the fly look quite alike externally, the maggots from the Vera- 
cruz specimen all show two large black bunches of strong more 
or less swollen and hooked or cleft spines on cephalic segment 
which are wholly lacking in the maggots from the Colorado 
specimen, indicating two very distinct species. The maggots 
from the Peruvian specimen do not show these cephalic spine- 
bunches; they besides differ from both the Colorado and Vera- 
cruz maggots in the characters of the segmental plates. 

The Veracruz specimen came from Orizaba (coll. by Herbert 
Osborn) and is probably P. bilimeki B. B., the type of which 
came from the same locality. A much fuller description with 
figures of the maggots and female reproductive system will be 
given in forthcoming papers. 

32. Megaprosopine series — Microphthalma spp. (North 
and South America) dissected, TD 313, 3915. Uterus pres- 
ent, long, subtubular, in several coils, filled with thousands of 
slender pointed eggs and maggots. The maggot of Microph- 
thalma is very hairy, being the extreme development in this 
respect so far as known. This series includes Microphthalma 
and allies, and almost certainly Megaprosopus. Trixodes is 
almost certainly the type of a separate series. The first and 
probably the second are white-grub parasites, while Trixodes is 
probably a woodboring-grub parasite. 

33. Macronychniine series — No dissections of Macrony- 
chia have as yet been made. The uterus may well be Meto- 
piine in form. At all events it must be quite distinct from the 
preceding series. The forms are perhaps muddauber-wasp 

34. CuTEREBRiNE series — Cuterebra spp. (Florida and 
South Carolina) dissected, eggs drawn, TD 487, 486. Probab- 
ly no uterus, as the eggs are certainly deposited ; chorion of egg 
very thick and hard, furnished at what is probably cephalic 
end and with a hinged lid or cap opening on dorsal aspect of 
egg and provided for the exit of the maggot which could not 
otherwise escape from its heavy chorion-prison, thus demon- 
strating most conclusively that the egg is intended for deposi- 
tion as such. Eggs probably deposited externally on skin or 

138 Annals Entomological Society of America [Vol. IV, 

hairs of host, and not swallowed, the maggots probably — almost 
certainly — penetrating skin at point of oviposition ; Dermatobia 
hominis is practically known to have these habits. Cuterebra, 
Dermatobia, and probably Rogenhofera and Bogeria. 

35. Gastrophiline series — Probably a uterus in which the 
eggs are incubated to a certain extent; eggs pediceled and per- 
haps ready to hatch soon after deposition, deposited on hairs 
of Equus and on Elephas, in the latter case probably on the 
hairs; maggots issuing from chorion in mouth or oesophagus of 
host and living in alimentary canal of same, passing out with 
the faeces when fully grown. Gastrophilus and probably 
Cobboldia, the latter parasitic in elephants in Africa and India. 

36. Oestrine series — Uterus certainly present; maggots 
deposited in nostrils of ruminants, elephant and horse, living 
in nasal and pharyngeal cavities and issuing through nostrils 
when ready to enter ground for pupation. Oestrus, Rhinoes- 
trus, Cephenomyia, Cephalomyia, Pharyngobolus — last known 
only as maggot in pharynx of elephant in Africa. 

37. Hypodermatine series — Peristomalia or facilia of adult 
fly remote, enclosing between them what seems to be the exces- 
sively broadened clypeus or epistoma, perhaps both differing 
in this character of the facial plate most widely from all the 
other Muscoidea. The sclerites of these parts need careful 
study before they can be rightly interpreted. Perhaps a uterus 
in which the eggs are partially incubated ; eggs deposited on hairs 
of ruminants, rodents and horse, taken into mouth of host, 
where they hatch or in oesophagus, the resulting maggots mak- 
ing their way slowly through the tissues to a position beneath 
the skin where they provide an air-hole and develop rapidly. 
Hypoderma, Oedemagena and Oestromyia. One of the most 
remarkable, widely divergent and at the same time well known 
groups of the superfamily Muscoidea. 

It is to be noted that the statements "dissected and drawn" 
and "dissected" refer always to the female reproductive and 
accessory organs; also that the term maggot as used refers 
always to the first-stage maggot only, unless otherwise specified. 
The term series above is not used in a strictly taxonomic sense. 
The maggots of most of the genera mentioned above for maggot 
characters have been drawn and described. 

Although much has been already accomplished in the inves- 
tigations above outlined, the results as tabulated most graphic- 

1911] Study of Muscoid Flies 139 

ally illustrate the astonishing amount of such work that yet 
remains to be done. The above 37 series, as defined or at least 
mentioned, and including others indicated therewith, will no 
doubt be more than doubled before the work is brought to a sat- 
isfactory conclusion. I have at present in tentative outline some 
91 groups— taxonomic categories which are to be considered of 
subfamily or tribal rank, perhaps better the latter, and which 
may be termed group units. But before these can be satis- 
factorily defined, hundreds of type species of genera must be 
carefully dissected and figured, as well for their eggs and mag- 
gots as for the reproductive system. It is now most confidently 
believed — in fact, it is axiomatically apparent — that this method 
of work will finally clear up the taxonomy of these flies and put 
it on a sound basis. 


In order to bring the outline of results up to date of reading 
proof of the foregoing (April 15, 1911), I wish to add the fol- 
lowing very brief particulars: 

Calliphorine series — Synthesiomyia belongs here, as shown 
by dissection of Peruvian material. It was formerly thought 
to be more closely allied with Musca. 

Mesembrinine series — Dr. Lutz has sent me three speci- 
mens in fluid of Pseudogametes for dissection, but unfortunately 
all are males. The fly has a wonderfully strong Oestrid habi- 
tus, greatly resembling Cuterebra in general form and appear- 
ance. It will almost certainly need a separate group, the 

Sarcophagine series — Sarcophaga auribarbata n. sp., auri- 
gena n. sp., argenta n. sp. (all Peru) have been dissected and 
drawn. Sarcophagula peruana n. sp. (Peru) and many other 
species of several genera (Florida and Peru) have been dissected 
and the maggots drawn. 

TD 354, from Massachusetts, which is apparently closely 
related to this group, if not a member of it, is remarkable as 
showing most clearly in the first-stage maggot the seven main 
pairs of sclerites of the cephalopharyngeal skeleton enumerated 
at the end of these addenda, with the single exception of the 
dorsopharyngeal sclerite which is rudimentary or nearly absent. 

140 Annals Entomological Society of America [Vol. IV, 

Metopiine series — Prof. Osborn's important observations on 
the habits of Senotainia (Ohio Nat., VII, 1906, p. 38) indicate 
that these flies are guided to the nests of the host by observing 
the latter in the act of transporting spiders or caterpillars with 
which to provision same. I believe the maggots are deposited 
in choria, and stuck to the spiders or caterpillars. After the 
nest is closed the increased temperature probably arouses the 
maggots to activity, whereupon they first devour the egg or 
grub of host and then attack the stored provision. The entire 
contents of the uterus, which are not great, are deposited almost 
certainly at one time, extending over several hours, and prob- 
ably may all be deposited in one nest if the conditions are fav- 
orable. Perhaps the fly attaches a maggot to the egg of host. 

CoMPSiLURiNE series — Eucelatoria australis n. sp. (Peru) 
has been dissected and drawn. The piercing larvipositor in 
this group is composed of two pieces, of which the upper is 
much shorter than the lower, the lower being concave dorsally 
and the upper concave ventrally. Both are sharply pointed 
but the lower piece curves broadly downward to the point and 
forms the main piercing organ. The two pieces are hinged at 
the posterior edge of their broad basal portion and the vagina 
opens between them. The maggot is expelled between these 
pieces within the skin of the host. Bouche, Heim and Kirsch- 
ner were right as to the piercing function of this organ, while 
Giard was wrong. Nielson adopted the latter's view in his 
criticism of my original announcement of subcutaneous larvi- 
position in these forms, to which I replied in Science (issue of 
Feby. 4, 1910, p. 195). 

There are other forms with what appears to be a piercing 
larvipositor. Mention has already been made of TD 480, 
which seems to be near Alophora (see Phasiine series). Another 
case is furnished in a fly from Massachusetts with pseudodexiine 
aspect, TD 371, which has a broad blade-like larvipositor. 
These, if true piercers, are probably cases of independent 
specialization not indicating relationship with this group. 
Emphanopteryx (not Cryptomeigenia) has a piercer-like organ 
in the female, but it is doubtful if it is functional as piercing 
the skin of the host. 

The genus Celatoria probably also possesses a piercing larvi- 
positor similar to that of Compsilura. Judging wholly from 
the descriptions and figures, Coquillett evidently misinter- 

1911] Study of Miiscoid Flies 141 

preted the sexes. The sex which he calls the male and which 
has the strong ventral keel is apparently the female, the keel 
being functional as a protection to the point of the larvipositor. 
The ventral groove described indicates this, and the fifth seg- 
ment mentioned is probably the broad basal part of the larvi- 
positor. If my surmise is correct, it appears that this form is 
unique in having the male front wider than that of female. 
At all events the genus must go in a group by itself, the Cela- 
TORiiNE, on the remarkable spinose character of the last-stage 
maggot and puparium combined with the other striking char- 

Gymnosomatine series — Gymnosoma sp. (California and 
Veracruz) dissected, TD 1815, 1815a. The uterus was not 
noted and is probably absent. The egg is large, white, very 
elongate-oval, flattened, chorion without reticulation. Gym- 
nosoma and allies, perhaps including Oedemasoma, Parasites of 
adult pentatomids. Habit, host-oviposition. 

Tachinine series — Neilsen is evidently wrong in his state- 
ment that his Tachina larvarum deposited maggots. The 
eggs which he referred to Carcelia were almost certainly those 
of his T. larvarum. This can be seen by reading his text. 
In his second paper (1910) he shows the egg of Carcelia to be 

Meigeniine series — This will need further division. The 
present forms are parasites of Chrysomelid larvae, sawfly 
larvae, and lepidopterous larvae. Tachinomyia appears to 
belong here. The following further groups can be indicated 
at present. 

ViviANiiNE series — For Viviania, Cryptomeigenia and allies. 
All are parasites of adult beetles so far as yet known. 

Thrixionine series — Thrixion and allies. Parasites of 
adult phasmids, etc. 

Plagiopine series. — Plagiops littoralis gen. et sp. nov. 
(Ocean Beach, So. Florida), TD 738, mentioned above under 
Meigeniine series, dissected. The uterus was not noted but is 
perhaps present. The egg is small, flattened, and ovate. 
Habit, host-oviposition. 

Plagiine ser — Siphoplagia sp. (Florida) dissected, TD 489; 
Cyrtophloeba sp. (Mass.), TD 2731, and Plagia sp. (Mass.) 
TD 2711, dissected. The uterus is in several coils and strap- 
like in upper extent, the elongate subcylindrical eggs packed in 

142 Annals Entomological Society of America [Vol. IV, 

rows and developing maggots therein. Habit, larviposition 
probably on host, as the uterine capacity is limited to a few 

Zygosturmiine series. — This is the 15th series above. 
Zygosturmia inca gen. et sp. nov. (Peru) dissected and drawn, 
TD 4031. Coquillett's Sturmia distincta and my protoparcis 
belong to this genus apparently. The oviducts are quite 
long and slender, common oviduct about two-thirds as long as 
oviduct, tubular glands about one and one-half times as long 
as common oviduct plus oviduct, spermathecal ducts about as 
long as common oviduct, tubular gland ducts only a little 
shorter than spermathecal ducts; ovaries with comparatively 
few ovarioles; no true preuterus; uterus strap-like, in but one or 
two coils, eggs and maggots packed in on the Echinomyiine 

AzYGOBOTHRiiNE series — Azygobothria aurea gen. et sp. 
nov. (Peru) dissected and drawn. The ovaries are large, of 
many ovarioles. Common oviduct is about as long as one 
ovary. There is a nipple-like preuterus. The spermathecal 
ducts are very long, fully twice as long as common oviduct, 
sinuate, bent at or below middle. Tubular glands are short, 
only half as long as common oviduct, their ducts nearly twice 
as long as the glands. The uterus is wide and strap-like, in 
about two coils more or less. 

Argyrophylax and Zygobothria seem to belong in the neigh- 
borhood of these two series, but may not come within either. 

Hystriciine series — The leaf-larvipositing forms will need 
division into at least seven groups, the present including 
Hystricia, Bombyliomyia, Jurinia and allies. The others 
follow : 

Melanophryonine series — Melanophrys and Artopharista 
which may or may not be the same. 

Panzeriine series — Panzeria, Varichaeta and their allies. 
Nielson has described the first-stage maggot of Panzeria as 
having no minute colored plates, which I think is a mistake. 
The fragment which he thought to be first stage of this species 
seems either not to have been this species or else to have been 
a piece of a second-stage skin. His text and figures indicate 
this conclusion. 

MiCROPALPiNE series — Micropalpus and allies. Cope- 
drypta may or may not come here. 

1911] Study of Muscoid Flies 143 

EcHiNOMYiiNE series — Echinomyia, Fabricia and allies. 
Peletria and their allies may come here. 

Saundersiine series — Saundersia, Epalpus and their allies. 

Dejeaniine series — Dejeania, Paradejeania and Lasio- 

Masiceratine series — This and the ISth and 19th series 
above represent only a part of the leaf-ovipositing minute-egg 
forms of Tachinidae, which exhibit a surprising variety of type, 
in female reproductive organs and eggs. They may be divided 
as follows, so far as now known, leaving the present group for 
Masicera, Blepharipa and allies, which have a long uterus con- 
taining black eggs with a honey-comb reticulation of the 

Brachymasiceratine series — This is the 18th series above. 
Brachymasicera polita gen. et sp. nov. (Peru) dissected, TD3987 
Short uterus, black eggs, chorion with a wrinkle pattern and 
not honey-comb reticulate. 

Belvosiine series — Belvosia, Latreillimyia and allies. Bel- 
vosia piurana sp. nov. (Peru) dissected, TD 4032. Uterus long, 
eggs black, chorion without reticulation but coarsely punctu- 
late, the punctures largely double and triple. 

Triachora has the chorion more finely and evenly j^unctu- 
late, the punctures all single. 

Blepharipezine series — Blepharipeza, Parachaeta and al- 
lies. Long uterus with black eggs, the chorion conspicuously 
honey-comb reticulate, the divisions with minute raised points. 

Goniine series — Many specimens of various spp. of Gonia 
from both North and South America have been dissected, and 
drawings made of two of the dissections which appear widely 
different. One is a short thick uterus in only one to two coils, 
with very short oviducts and large ovaries, TD 4011; the other 
is a very long uterus in five or more coils, with elongate oviducts 
and somewhat smaller ovaries, TD 4037. Both are from Peru. 
Other specimens from Piura, Nana (near Lima), and Arequipa, 
Peru, show both long and short uteri, and I have found developed 
maggots in the short uterus as well as in the long one. The eggs 
are minute, flattened ventrally, ovate to subcircular; the chorion 
is without reticulation, but evenly, closely and very finely 
punctulate throughout. Preuterus present, common oviduct 
short, spermathecal ducts and tubular glands elongate. The 
maggot is elongate-pyriform, sometimes appearing nearly sub- 

144 Annals Entomological Society of America [Vol. IV, 

cylindrical, and appears to lie curled in the sub-circular or oval 
egg. Pressure on the egg often causes it to assume an elongate 
narrow form pointed at ends, inside which the maggot probably 
lies straight. Slight pressure or teasing with a needle causes 
the cylindrical anterior portion of the maggot to protrude from 
the ventral edge of egg, at right angle to long axis. I can see no 
difference in the maggots from the short and long uteri, and am 
constrained to believe that here the uterus gradually increases 
in length as the eggs descend from the ovaries and fill it, although 
oviposition may begin while the uterus is still short. If this is 
so, it is an exceptional case in the great increase of length after 
maggots have become fully formed within the choria. There 
seem to be two forms of the flies in the Peruvian material, dis- 
tinguishable externally, but the separation does not accord with 
the uterine difference. One or both of these forms are prob- 
ably what has been called Gonia chilensis. 

My dissections have shown Gonia frontosa (Mass.) to have 
a long coiled uterus; and what I identify as G. pallens and G. 
angusta (S. Florida) to have a very long uterus. The chorion 
of the egg is the same in these forms as above described. 

The above observations indicate that much care and judg- 
ment must be used in describing the female reproductive organs 
in these flies. I have noted variations in lengths of uterus 
in other forms within certain limits; and variation in the number 
of rows of eggs in the strap-like forms of uterus as well as in 
certain others, without a corresponding variation in length. 
It is evident that we need careful investigations in various 
groups which shall determine how many days elapse between 
issuance of the fly or fertilization and complete development of 
the female reproductive organs. I have noted that in Vari- 
chaeta, under artificial conditions, this takes about fourteen 

Phasiatactine series — Phasiatacta elongata gen. et sp. 
nov. (Peru) dissected and drawn, TD 4019. Uterus extremely 
long and slender, in very many coils and irregular turns, preu- 
terus present. Ovaries large, oviducts long and slender, sper- 
mathecal ducts long and curved, tubular glands elongate. Eggs 
black, elongate, pointed at each end, chorion with areoles gath- 
ered around a dorsal opaque area, without reticulation. 

Cnephalomyiine series — Cnephalomyia floridana gen. et 
sp. nov. (So. Fla.) dissected, TD 877. Uterus very long and 

1911] Study of Muscoid Flies 145 

slender, eggs blackish, elongate, obtusely pointed at each end, 
chorion, with an almost perfect hexagonal reticulation, inter- 
spersed with fine punctulations. 

Cnephalodes pollinosus gen. et sp. nov. (Peru), TD 4038, 
which has been dissected and drawn, probably belongs here. 
The uterus is extremely long and slender, in numerous irregular 
coils and turns, the eggs elongate in single to quadruple file, 
chorion black with honeycomb reticulation and punctures. 
The characters of the maggot, including those of the cephalo- 
pharyngeal skeleton, show a close relationship with Gonia. 

Ophirosturmiine series — This is the 19th series above. 
Uterus long and in many coils, eggs brownish-yellow, chorion 
honey-comb reticulate interspersed with closely-set microscopic 
raised points. Oviducts long, spermathecal ducts very long 
and slender, tubular glands moderately long, preuterus present. 
Phasmophaga evidentally does not belong here. 

Ommasiceratine series. Ommasicera chaetosa gen, et sp. 
nov. (Peru) dissected and drawn, TD 4018. Uterus short and 
thick, in only one and a half or two coils ; eggs brownish-yellow, 
chorion with an elongate-hexagon reticulation like a honey- 
comb pattern drawn out of shape longitudinally. 

This concludes the leaf-ovipositing groups so far as I am 
able at present to indicate them. But there remains a multi- 
tude of these forms yet to be assigned, among which I can give 
notes on the following: 

Paragermaria has a very long slender uterus, about 40 mm. 
in length, eggs black, chorion without reticulation, finely punc- 
tulate much like that of Triachora. It probably goes in the 
Germariine series, which will include Germaria and allies. 

Cnephalia has a very long and slender uterus containing 
black eggs. 

Attacta, as represented by a specimen from Guatemala 
which I doubtfully refer to this genus, has black eggs whose 
chorion is not honey-comb reticulate but shows minute punc- 
tures in a roughened surface. The uterus is probably elongate. 
The puncture pattern of the chorion appears in the peripheral 
area like a chitinous network, with an elongate or oval less- 
closely punctured central area. The surface of the chorion is 
closely and minutely roughened with projecting points. 

Ceromasia has brownish-yellow eggs, the chorion is intensely 
honey-comb reticulate, each division closely set with micro- 
scopic raised points. 

146 Annals Entomological Society of America [Vol. IV, 

Phasmophaga has a brownish-yellow egg, the chorion appears 
not to possess any reticulation and is not punctulate; the sur- 
face appears to be quite smooth. 

TD 390 (Mass.) is a small fly with a disproportionately long 
uterus; it has much the habitus of Eusisyropa, and has been 
determined by Mr. W. R. Thompson as Masicera sp. near 
pauciseta. The egg is deeply yellow with a slight brownish tinge, 
the chorion is reticulate with nearly perfect hexagons, and the 
surface is sparsely set with long chitinous spine-like points which 
look like short hairs growing therefrom. No punctulation. 

TD 355 (Mass.) was at first determined by me as Eusisyropa 
blanda and so published in Tech. ser. Bull. 12 Bur. Ent., p. 
116. I doubt if it is that genus. It contained some 700 whitish 
eggs in the uterus, which was thus probably of the short thick 
type. These eggs show the nearly mature maggots within. 
The chorion is thin, without reticulation or punctulation, and 
has much the appearance of the Phasmophaga chorion exter- 

Finally, TD 747, a small yellow-legged fly with a pseudo- 
dexiine habitus and a very long slender uterus, from Ocean 
Beach, South Florida, shows black eggs whose chorion appears 
very similar to that of Phasmophaga in structure. There is no 
punctulation or reticulation, and practically no appearance of 
rugosity, the surface being quite smooth. Filaments of trans- 
lucent substance appear to depend from the ventral surface 
of this egg, showing in every one of man}^ mounted specimens 
as pendulous loops and coils. I have seen nothing of the kind 
in any other egg. They are perhaps intended for more secure 
attachment to leaf surfaces. 

These examples might be multiplied, but I have given suffi- 
cient to show the wonderful diversity of type exhibited by these 
minute-egg forms, all of which are believed to oviposit upon the 
foliage of the plants. 

Ophirionine series — This name should be applied to the 
21st series above. 

EuMYOBiiNE series — Eumyobia flava gen. et sp. nov. (Peru) 
dissected and drawn, TD 4021. Ovaries large, oviducts short, 
common oviduct long, spermathecal ducts and tubular glands 
short, preuterus present; uterus very heavy and thick, in two 
stout coils or so, the elongate maggots developing therein. 
The eggs and maggots are packed in irregularly, in various posi- 

lull] study of Miiscoid Flies 147 

tions. Pyrrhosia may belong here, and the present form may be 
that genus or one of its near allies, being apparently what Coquil- 
lett determines as Leskia. I have no alternative, however, but 
to name it to prevent doubt hereafter. 

PsEUDODEXiiNE series — There are almost certainly many dis- 
tinct groups here. The Orphirodexiine series must be erected 
for Ophirodexia and its allies. 

Atrophopodixe series — This is quite distinct from Ophiro- 
dexia, as shown by Diaphoropeza peruana sp. nov. (Peru) dis- 
sected and drawn, TD 4026. This species is not typical of the 
genus but comes nearer to it than to any other, and certainly 
belongs to the Atrophopoda group. Ovaries not large, oviducts 
very short, common oviduct very long, spermathecal ducts 
and tubular glands short, preuterus present; uterus thick and 
swollen, in one coil or so, eggs and maggots very oblique to 
longitudinal. It is as yet doubtful if Vanderwulpia can be here 
included. The group is proposed for Atrophopoda and its 

Beskiine series. — Beskia and ahies. No proper material 
for dissection has yet been available but the group must be 
pointed out as probably furnishing a most marked deviation in 
larval habit from any hitherto known in the muscoid flies. 
In 1908, Mr. E. O. G. Kelley found, at Pawnee, Oklahoma, a 
maggot which he observed at the time to be, to quote from Mr. 
Webster's letter on the subject, "in the act of actually feeding 
on a grain aphid of wheat, probably Macrosiphum granaria." 
This maggot changed to a puparium which was attached to the 
base of a wheat leaf. From this puparium issued a fly of Ocyp- 
terosipho aelops, which is very close to Beskia if not the same. 
No confirmation of this larval habit has come to light since, but 
although the record is a most surprising one it is probable that 
the maggots of these flies are external feeders on aphids and per- 
haps some other soft-bodied insects. The anal stigmata of the 
puparium are borne at the ends of long processes which suggests 
an analogy with aphidophagous syrphids. 

Steinielline series — From what Nielsen has shown of the 
characters of the first-stage maggot of Steiniella, the genus repre- 
sents a separate group. This maggot possesses minute colored 
subchitinized plates interspersed with short spines, some of the 
plates bearing spines on their posterior edge. The maggots are 
almost certainly not deposited on leaves. The form is not 

148 Annals Entomological Society of America [Vol. IV, 

allied with the Hj^striciine group of series, the adult being very 
distinct in external characters. The minute colored plates of 
the first stage maggot are evidently an independent specializa- 
tion, and are interesting as indicating the probable method of 
development of the more perfect colored scale-like plates of the 
leaf-larvipositing forms. 

PsEUDOMYOTHYRiiNE series — This is the 27th series above. 
Pseudomothryia perplexa sp. nov. (Peru) dissected and drawn, 
TD 4001, 4035. Ovaries small, of only about six ovarioles. 
Oviducts about one-third as long as ovar3^ common oviduct 
about twice as long as oviduct, spermathecal ducts very elon- 
gate, tubular glands about the length of latter, preuterus 
absent. Uterus slender, laterally compressed, in three to four 
coils, inside each other disposed in a more or less regular spiral, 
eggs and maggots in single file. The oral spine which terminates 
the mandibular sclerite of the cephalopharyngeal skeleton in the 
first stage maggot is very sharp and needle like. Methypostena 
and Tachinophyto perhaps come here. It is quite likely that 
Erynnia nitida, the European tachinid parasite of the imported 
elm-leaf beetle, occurs in North America and had been deter- 
mined by Coquillett as Hypostena barbata. 

EuzENiLLiiNE Series — Euzenillia aurea gen. et sp. nov. 
(Mass.) dissected, TD 350. In this dissection the reproductive 
organs were not extracted intact, and I can judge only from the 
maggot which is a wholly new type. The latter is elongate, 
moderately slender, and thickly clothed all over with short 
fine minute spines except on posterior half of dorsum. The 
cephalopharyngeal skeleton is rather slender. The spermathecae 
are suboval or elongate-rounded, and the larvipositor is normal. 
A uterus is present, which is probably not long. 

Parexoristine series — Carcelia belongs here. Nielson, in 
his second paper (1910), has shown the egg to be elongate and 
pediceled, as in Parexorista. 

Dexiine series — This series needs splitting into several 
groups. Aside from the typical group, to include Dexia and 
allies, the following may be given as at present recognizable: 

Almugmyiine series — For Almugmyia and allies, whose 
reproductive characters are mentioned under the Dexiine series 
above. Microchaetina probably comes here, and may prove 
to be the same as the present form. 

1911] Study of Muscoid Flies 149 

SiROSTOMiNE series — This will include Sirostoma and its 
allies, which are white-grub parasites. The females deposit 
their maggots at the surface of the soil, into which the latter 
penetrate in search of hosts. Phorostoma probably comes here. 

Sardioceratine series — Sardiocera and allies. Theresia and 
Eutheresia gen. nov. for Coquillett's Theresia analis, probably 
come here. All are parasites of wood-boring grubs of Coleop- 
tera. The host relations of Eutheresia are most interesting, 
and will be detailed elsewhere. The maggots of Sardiocera 
(which I determine as Coquillett's Theresia tandrec) and 
Eutheresia both have a pair of short anal processes carrying the 
tracheas with the anal stigmata at their ends. 

Prosenine series — For Prosena, Myiocera and allies. 

EcHiNODEXiiNE series — For Echinodexia and allies. Prob- 
ably Hystrichodexia, Eudexia, Hystrisiphona, and Bathydexia 
come here. These forms are distingiushed in the fly by having 
spine-like macrochaetae on scutellum and abdomen, and would 
thus seem to form a natural group by themselves. This char- 
acter probably does not carry with it the leaf-larviposition 
habit, although the flies bear a strong superficial resemblance 
to the Hystriciine and allied forms. It will be noted that spine- 
like macrochaetae occur in the Blepharipezine series of the leaf- 
ovipositing minute-egg forms; also that this character does not 
extend to all the leaf-larvipositing forms. 

Myiophasiine series — The females of Ennyomma globosa, 
parasitic in Chalcodermus aeneus in cowpea pods, probably 
deposit the living maggots at the weevil punctures of a certain 
age, and the very slender maggot bores in next the periphery 
of the hardened-sap pellicle which closes the puncture until it 
reaches the semiliquid frass, through which it can easily gain 
access to the host grub. 

Phasiopterygine series — The peruvian species mentioned 
above, TD 4005, is Phasiopteryx australis sp. nov. It has been 
dissected and drawn. The uterus is about sixty millimeters in 
length, from one-sixth to one-half millimeter in width, and is 
perhaps the extreme development for relative length and slen- 
derness in the superfamily, though Phasiotacta is a close second. 
On page 78, T. A. E. S., XIII (March, 1895), I mentioned two 
specimens of a fly from Doctor Forbes, Ills., as doubtfully refer- 
able to Ormia, one of which was reared from Crambus sp. If 
these specimens belong to the Oestrophasiine series, as is quite 

150 Annals Entomological Society of America [Vol. IV, 

probable, this will form the only rearing record yet known of 
any immediate relative of this group. Here seems to be a hint 
explanatory of the remarkable specialization of the first -stage 
maggot in Phasiopteryx, for it is probable that Oestrophasia has 
a similar maggot. The larvae of Crambus work underground, in 
silk-hned galleries, feeding on the stems of growing plants, 
especially corn. The Phasiopteryx type of maggot would be 
quite well adapted to search out such hosts. Its smooth seg- 
mental plates would enable it to penetrate the silken walls of 
the galleries, where an ordinary maggot would become enmeshed. 
Yet one cannot help thinking that the Phasiopteryx maggot is 
rather needlessly specialized for such prosaic purpose. Per- 
haps Oestrophasia has a still difTerent type of maggot. 

Characterizations to fit the new generic and specific names 
used in this article will be shortly published, probably in the 
more exhaustive paper from which the whole of the present 
data are taken in advance. I must explain that I consider it 
quite obligatory upon me, in all cases of doubt as to the deter- 
mination, to give new names generic and specific to the forms 
dissected, described and figured. These names will fix the 
forms so as to prevent any further doubt arising as to their 
identity. If this plan increase the synonymy, as it doubtless 
will to an extent, there is no positive harm done; while the posi- 
tive good is secured of certainty in determination. 

In the forthcoming more complete paper will also be given 
fuller explanations of the functions of the various parts of the 
female reproductive system ; the necessity for the use of the terms 
preuterus, uterovagina, etc., which may be thought by some to 
be superfluous, with numerous plates illustrating all the repro- 
ductive types that have been drawn. 

I wish to announce that, after an extended comparative 
study of many maggot types. I find the first-stage cephalo- 
pharyngeal skeleton to consist of seven original main sclerites, 
all paired, as follows: 

1. Pharyngeal — "lateral pharyngeal" of Hewitt in whole or part; "upper 

pharyngeal" of Nielsen plus "lower pharyngeal" 
of same author in whole or part. 

2. Infrapharvnoeal — lower wing of "lateral pharyngeal" of Hewitt in 

part; "lower pharyngeal" of Nielson in part. 

3. DoRSOPHARYNGEAL — "dorsal pharyngeal" of Hewitt. 

4. Hypostomal — after Hewitt. 

5. Infrahypostomal — "the plate below the skeleton "of Nielsen. 

6. Mandibular — after Hewitt. 

7. Dentate — after Hewitt. 

1911] Study of Muscoid Flies 151 

A considerable number of minor sclerites occur in various 
forms, but all seem to be derived from the above seven main 
ones, except those of the oral region which belong to the pseudo- 
cephalon and are probably developed from it. I have an ex- 
tensive series of drawings of a very large number of types of 
first-stage skeleton, which will be published in due time in 
connection with a comparative study of all the sclerites that go 
to make up the skeleton in the various forms. 

Under the Sarcophagine series in these addenda is mentioned 
TD 354 from Massachusetts, which most clearly shows all the 
above main sclerites except the dorsopharyngeal. The infra- 
pharyngeal is so clearly exhibited in this form as to prove beyond 
doubt, I think, its distinctness as one of the main sclerites. 
In most forms the infrapharyngeal, while more or less present, 
is so intimately connected or welded with the pharyngeal as to 
appear a part of the latter. This has misled Nielsen and others. 

It is worth while mentioning in a speculative way that the 
above seven main pairs of sclerites may represent the seven 
embryonic segments absent in the muscoid maggot, which have 
been inverted to form the skeleton proper of the maggot mouth 
and pharynx. The head in the most primitive insects is believed 
to be composed of seven segments and the abdomen of ten, the 
three thoracic segments bringing the total number to twenty. 
There are twelve segments in the muscoid maggot besides the 
pseudocephalon. Dr. C. Gordon Hewitt's admirable mono- 
graph of Musca may be studied with much profit in this con- 

What I have heretofore called the clypeus in the fly 
(Taxonomy, pp. 22-24) seems to be a part of the everted dorsal 
wall of the pharynx chitinized. The true clypeus is apparently 
the lower portion of what I have called the facial plate. The 
labrum seems to have projected itself forward, carrying with it 
the epipharnyx to form the dorsal part of the haustellum, and 
in this way becoming widely separated from the clypeus. 
Into the space thus left there appears to have been everted the 
chitinous portion of the pharynx to form the rostrum of the 
proboscis, which attaches basally in front to the epistoma or 
anterior edge of the clypeus. The morphological sequence of 
the parts is thus lost when the proboscis is extended, but when 
the latter is retracted the rostrum is inverted to a semblance of 
its original morphological position. 

152 Annals Entomological Society of America [Vol. IV, 

It should be stated that the ultimate categories of genera, 
or those which will eventually be found most serviceable for 
taxonomic recognition and which I shall call group units, 
may be considered as generally equivalent to tribes in other 
groups of insects. Most of the series given in these addenda 
are to be considered as such, and for that reason I have given 
them the names of the genera dissected and studied. 

The article by me in a recent issue of "Science" entitled 
"On muscoid and especially tachinid synonymy, may be pro- 
fitably read in connection with this preliminary outline of results- 
to date in the Muscoidea. 


James S. Hike. 

The species of these two genera from America north of 
Mexico, have been considered by WilHston in Volume XII, of 
the Transactions of the American Entomological Society, and 
the Mexican species of Promachus have been tabulated by 
Osten Sacken in Biologia, Diptera I, 192. The fact that it has 
been possible to get together an extensive collection of spec- 
imens of various species from different parts of North America 
has led me to make an attempt to identify many of the Nearctic 
forms, and as some success at least has been attained the fol- 
lowing results are offered for the consideration of those, who 
for any reasons, have interest in the various species included. 

One can not study these flies in the field without noting 
their highly predaceous habits. These habits have attracted 
the attention of observers in the past with the result that a few 
of the species have been mentioned as injurious from the stand- 
point of killing honey-bees and other useful insects, but in most 
cases the feeding habits are variable to the extent that each 
species accepts a large variety of insect life as food, so that 
after all it seldom happens that their attacks are concentrated in 
a particular direction sufficiently to establish a marked habit 
which shall be considered either injurious or beneficial. They 
are among the largest species of their subfamily, some speci- 
mens measuring nearly forty-five millimeters in length and the 
smallest over twenty millimeters, so if their predaceous habits 
could be controlled the results, very likely, would be of extraor- 
dinary importance. 

The material used in the preparation of this paper has been 
procured from various sources and I am under many obligations 
for favors. The United States National Museum and the 
museums of Cornell University and the University of Kansas 
each have loaned specimens for study. Professor J. R. Watson, 
of Albuquerque, New Mexico, has sent me several specimens 
from that interesting region; Professor J. H. Schafifner has 
donated the material he collected during two summers in cen- 
tral Kansas; Professor C. F. Baker has sent me much material 
that D. L. Crawford collected in Mexico during the season of 
1910; while Charles Dury of Cincinnati, and H. S. Harbeck, of 
Philadelphia, and others have forwarded various species from 


154 Annals Entomological Society of America [Vol. IV, 

their respective localities. D. W. Coqidllett, Chas. W. John- 
son, J. M. Aldrich and other Dipterologists have aided in vari- 
ous ways. One finds it much pleasanter to work at a difficult 
task when he meets with such encouragement as I have had in 
this study. 

The characters available for the separation of the species in 
l)oth of these genera are rather meager and I doubt not that 
students will experience more or less difficulty in using the keys 
and descriptions. The difi^erences which distinguish species 
are sufficiently evident, however, for the purpose when the 
proper comparisons can be made, but without the opportunity 
to make comparisons as is the case when a single species is 
considered one may not always feel satisfied with results. 

I was not able to get all of the described species from Mex- 
ico and Central America but where I possessed specimens of 
species from these countries they are included in the consid- 
eration of their respective genera. 

A list of the species and synonymy is given lielow but the 
reader is referred to Aldrich's Catalogue of North American 
Diptera and to other works for further references to literature. 

List of the Species or Proctacanthus. 
arno Townsend, Proc. Cal. Acad. Sci. IV, 599. 

brevipennis Wiedemann, Auss. zw., I, 431. Van dcr Wulp, Tijdsch. 
v. Ent., XXV, lOS. Williston, Trans. Amer. Ent. Soc. XII, 73. 
caudatus new species. 
coquillettii new species. 
duryi new species. 
fulviventris Macquart, Dipt. E.xot., vSuppl. IV, SS. Osten Sacken, Cat- 

aloj^ue 23.), note. 
heros Wiedemann, Auss. zw., I, 427. Schiner, Verb. Zool. Bot. Ges., 

1S()(), (),S2; 1S(J7, 39(j. Willi.ston, Trans. Amer. Ent. Soc, XII, 74. 
longus Wiedemann, Dipt. Exot., 1S3; Auss. zw., I, 42(). Macquart, 

Hist. Natur. Dipt., I, 307; Dipt. Exot., I, 2, 123. Schiner, Verb. 

Zool. Bot. Ges., 1.S6G, ()<S2; 1867, 396. 
craverii Bellardi, Saggio, II, .50. Williston, Biologia, Dipt., I, 327. 
micans Schiner, Verb. Zool. Bot. Ges., 1867, 397. 
milbertii Macquart, Dipt. Exot., I, 2, 124. 

missouriensis Riley, 2d Mo. Report, 122, fig. 89. 
Pagrion Jacnnicke, Neue Exot. Dipt., .'57. 
nigriventris Macquart, Dipt. E.xot., I, 2, 124. 
nigrofemoratus new species. 
occidentalis new species. 
philadelphicus Macquart, Dipt. Exot., I, 2, 123. Williston, Trans. Am. 

Ent. Sue, XII, 7.). 
rufiventris Macquart, Dipt. Exot., I, 2, 123, plate X, figure 2. 
rufus Williston, Trans. Am. Ent. Soc, XII, 74. 

1911] Proniachus and Proctacanthiis 155 

Of the other North American species that have been placed 
in Proctacanthus, exquisitus Osten Sacken must belong to the 
genus Asilus if one may judge from the figure, for the posterior 
branch of the third vein meets the margin of the wing dis- 
tinctly behind its apex; P. virginianus Van der Wulp has the 
appearance of a species of Erax. The specimen figured is a 
fem.ale and no spines are shown at the tip of the ovipositor; 
this fact, together with the presence of a stump of a vein at the 
base of the anterior branch of the third longitudinal suggests 
Erax, for I have never found a species of Proctacanthus with 
this character present; Proctacanthus zamon Townsend has 
been known as a synonym of Eccritosia amjjhinome Walker for 
some time. 

List of the Species of Promachus. 
albifacies Williston, Trans. Am. Ent. Sec, XII, (53. Osten vSacken, 

Biologia, Diptera, I, 195. 
bastardii Macquart, Dipt. Exot., I, 2, 104. 
laevinus Walker, List II, 108. 
rtibi^inis Walker, Dipt. Saund., 123. 
ultimus Walker, Dipt. .Saund., 13(). 

philadelphicus Schiner, Vcrh. Zool. Bot. Ges., 1SG7, 389. 
fitchii Osten Sacken, Catalogue, note 121, 234. 

apivora P'itch, Country Gentleman, XXIV, 03; 3d N. Y. Report, 
251, plate IV, figure 7. 
forfex Osten Sacken, Biologia, Diptera, I, 194. 

quadratus Bellardi, Saggio, II, 27, plate II, figure 3. 
giganteus new species. 
minusculus new species. 
nigrans new species. 
nigripes new species. 

princeps Williston, Trans. Am. Ent. Soc, XII, 02. 
quadratus Wiedemann, Dipt. E.xot., 201; Auss. zw., I, 4S5. Osten 

Sacken, Biologia, Dipt., I, 192. 
rufipes Fabricius, Syst. Ant., 794; Syst. Antl., 109. Wiedemann, Dijjt. 
Exot., 203; Auss. zw. I, 4.S7. Williston, Trans. Am. Ent. Soc, 
XII, 107. 
sackeni new species, 
truquii Bellardi, Saggio II, SO, plate II, figure (i. W'illiston, Biologia, 

Dipt., I, 322. 
vertebratus Say, Journ. Acad. Sci. Phil., Ill, 47; Compl. Works, II, 02. 
Williston, Trans. Am. Ent. Soc. XII, 02. 

Five species of Promachus from Mexico and Central Amer- 
ica could not be procured so they are not considreed in this 
paper. They are anceps Osten Sacken, cinctus Bellardi, mag- 
nus Bellardi, nobilis Osten Sacken and pulchellus Bellardi. 

156 Annals Entomological Society of America [Vol. IV, 

Promachus fuscipennis, according to Osten Sacken, is not a 
North American species and Promachus trapezoidalis Bellardi 
is rightly included in the genus Mallophora as Williston has 
stated for it has the claws distinctly blunt at the tips. It may 
be of interest to state that this latter species has been taken at 
Brownsville, Texas. Heretofore it was known only from Mexico. 


Proctacanthus was described by Macquart in 1838. The 
species falling here are all rather large and robust, the marginal 
cell is closed, palpi one segmented, antennal bristle bare, veins 
closing the discal and fourth posterior cells not parallel, the 
posterior branch of the third vein curves forward to meet the 
costa before the tip of the wing, anterior branch of the third 
vein not angulated at base nor does it bear a stump, ovipositor 
cylindrical and with a terminal circlet of spines which may be 
very small in some species, abdomen longer than the wings. 
These characters are all well marked and there is no difficulty 
in placing the various species in the genus. The venation is 
much like Erax but the circlet of spines at the tip of the ovi- 
positor and the lack of silver white segments at the posterior 
part of the male abdomen are distinctive. 


1. Segments of the abdomen thickly pilose on the sides, very large red species. 

Abdomen pollinose or sparsely hairy, smaller species 2 

2. Femora black, tibiae red, the colors strongly contrasting 3 

Femora and tibia of nearly the same color, so there is no strong contrast in 

colors 4 

■3. From above the hypopygium is wide, and longer than segments seven and 
eight combined. Male abdomen largely red, female abdomen not red. 

Hypopygium small, distinctly shorter than segments seven and eight com- 
bined. Abdomen of both sexes largely red above fulviventris 

4. Abdomen red, except the first segment and anterior border of the second. . . .5 
Abdomen in large part gray pollinose ■ 6 

5. Dorsum of the thorax uniformly brown, hypopygium from above wide, appen- 

dages longer than segments seven and eight combined rufus 

Dorsum of the thorax usually striped with black, hypopygium small, the 
appendages from above shorter than segments seven and eight com- 
bined rufiventris 

■6. End lamella of the ovipositor densely pilose, spines at the tip small 7 

End lamella of the ovipositor sparsely hairy and with a circlet of enlarged 

spines at the tip 8 

7. Wings nearly hyaline, many of the veins faintly margined with brownish. . . . 

Rather small species, middle of the wing with a large dark colored patch 


.8. Wings hyaline or with only the veins margined with brown !) 

Wings of a uniform brown all over 12 

1911] Promachus and Proctacanthus 157 

9. Scutellum with white bristles duryi 

Scutellum with black bristles 10 

10. Wings clear hyaline, all the femora entirely black nigrofemoratus 

Wings with the veins margined with brown, at least the posterior sides of 

all, the femora red 11 

11. Male genitalia elongate, the appendages curved inward at the tips and enclos- 

ing an open space beyond the other parts arno 

Male genitalia short, appendages but slightly longer than the other parts 


12. Thorax with brown markings, wings clear brown 13 

Thorax with dark, nearly black, markings, wings pale brown 14 

13. Wings long and wide, palpi with white hair, two or three bristles on the 

front side of the hind femora besides those near the apex longus 

Wings rather short and narrow, palpi with black hair, four or five bristles 
on the front side of the hind tibia besides those near the apex 


14. Abdomen very dark, nearly black pollinose nigriventris 

Abdomen lighter, gray pollinose 15 

15. Large stout species, middorsal stripe of the thorax gray pollinose and not 

plainly differentiated milbertii 

Small, rather slender species of southern distribution, middorsal thoracic 

stripe usually plainly differentiated and not distinctly pollinose 


Proctacanthus heros Wiedemann. 

Total length, 32 to 45 millimeters. Body red all over, mystax and 
beard straw yellow, occipito-orbital bristles straw yellow; legs red with 
black bristles and yellow hair; bristles of the scutellum black; wings 
brown,' some of the cells along the inner margin nearly hyaline at the 
middle. First four or five abdominal segments on each side with an 
area of dense yellow pile, hairs on the dorsum of these segments and on 
all parts of the posterior segments short and black. 

The large size of this species together with the red color of 
the entire body and the pilose areas on the sides of the abdomen 
easily characterize it. 

Specimens from Southern Pines, North Carolina, collected 
by A. H. Manee. Also others from Georgia and Mississippi. 

Other writers have reported it from Florida, South Carolina 
and Kentucky. 

Proctacanthus rufus Williston. 

Total length 30 to 36 millimeters. Body red, with the exception of 
the first abdominal segment and the anterior margin of the second 
which are black. Mystax and beard yellow, the latter lighter than the 
former, palpi red with pale hair; thorax and legs mostly with dark hair 
and bristles, scutellum with numerous black bristles, wings nearly 
uniformly pale brownish, dorsum of the thorax plain red without distinct 
markings of any kind. Abdomen black at base above, otherwise red, 
the former color extends back to the anterior third of the second seg- 
ment. Male genitalia enlarged, wider than the last abdominal seg- 
ment, appendages from dorsal view wide at the base and narrowed 
towards the apex. 

158 Annals Entomological Society of America [Vol. IV, 

The red legs, uniform red thorax and enlarged hypopygium 
are characteristic. 

Specimens are at hand from Ohio, Arkansas, Kentucky, 
New Mexico and Massachusetts. Others record it from 
North Carolina and New Jersey in addition. 

The species appears to be partial to bare sandy areas such 
as are present along lakes, seas and rivers and in semi-arid 
regions. The larval and pupal stages are passed beneath the 
surface of the ground, and the pupa case is left partially pro- 
truding when the adult issues. 

Proctacanthus rufiventris Macquart. 

Total length 30 to o.j millimeters. This species is very much like 
rufus in coloration. In fact the only noticeable difference between the 
two is to be found in the markings of the thorax and the male genitalia. 
In most specimens of rufi\'cntris the thorax is marked with dark, nearly 
Ijlack stripes and spots. The hypopygium is small, narrower than the 
last abdominal segment and the appendages from dorsal view are 
narrow for their entire length, short and rounded at the tips. 

Specimens at hand from Louisiana, North Carolina, and 
Texas. Others record it from San Domingo, Honduras, and 
Porto Rico. 

Proctacanthus fulviventris Macquart. 

Total length 2.5 to 30 millimeters. Mystax and beard bright yel- 
low, palpi black with yello\\- hair, proboscis and antennae black. Thorax 
black, scutellum with numerous yellow bristles and hairs; wings pale 
Ijrown, femora clear black, tibiae red, except the apex of each which 
ma>^ be black, tarsi black or in some specimens the hind metatarsi are red. 
Abdomen black at the base, partly red posteriorly. Different individuals 
are colored differently. In two specimens before me the female abdo- 
men is clear black except the dorsum of segments three to seven inclu- 
sive which is red, while in the male the segments beyond four are red, 
as is the dorsum of three and four, while the venter of the first four seg- 
ments and the dorsum of the first two are practically all black. Male 
genitalia small, appendages much shorter than the last two abdominal 
segments, each widest at the Ijase and gradually narrowed towards the 
apex which is rounded. 

The small male genitalia together with the clear black 
femora and red tibia characterize the species. 

The specimens before me are from Florida and records do not 
show that it has been collected in any other locality. 

1911] Peomachus and Proctacanthiis 159 

Proctacanthus caudatus n. sp. 

Total length 2S to 30 millimeters. Mystax and beard pale yellow, 
palpi yellowish red with pale yellow hair, antennae and proboscis black; 
thorax dark with some light pollinose markings, bristles of thorax 
black, numerous but not very stout black bristles on the scutellum; 
femora black, tibia red, black at the apices, tarsi black; wings pale 
brownish. Male abdomen yellow, except -first segtnent and the base of 
the second which are largely black, hypopygium from dorsal view wide, 
from side view widened near the apex which is obliquely truncate. 
Female abdomen entirely dark, except the posterior margin of each of 
the second to fifth segments which are red; first abdominal segment in 
both sexes with numerous yellow hairs on each side. 

The male type and one female from \'eracruz, Mexico, col- 
lected by D. L. Crawford, of Pomona College, Ontario, Cal. 

The male hypopygium is very different from that of fulvi- 
ventris or rufus. 

In Volume I, page 206, Biol. Cent. Am. Osten Sacken men- 
tions a female of a species of Proctacanthus which appears to 
be the same as this one. His specimen was taken at San 
Geronimo, Guatemala. 

Proctacanthus occidentalis n. sp. 

Total length 30 to 36 millimeters. A modest colored species ha\'ing 
nearly clear wings with many of the veins narrowly margined with 
brown. Mystax very pale yellow, usually with a few black bristles, 
beard white, pal])i black with some black and some white hair, occipito- 
orbital bristles partially white and partially black; thorax pale brown- 
ish or gray, mostly with black bristles, scutellum with numerous black 
bristles and scattering fine white hairs, humeral callus distinctly lighter 
gray than the other parts of the thorax and toward the median line 
from each of these is a small white spot which is of some use in deter- 
mination for most other species do not have such a marking. Wings 
nearly hyaline, in most specimens the longitudinal veins are faintly 
margined with brown; femora red posteriorly and nearly black anteri- 
orly, tibiae colored with reddish and blackish in \'arious combinations; 
abdomen dark, gray pollinose, male genitalia shorter than the last two 
abdominal segments, appendages curved towards each other at the tips 
so that they nearly meet; female genitalia with the end piece densely 
yellowish pilose and with the spines at the tip so small that they are 
not plainly differentiated from the other hairs of the region. 

Male type from Los Angeles County, California, and eleven 
other specimens from California and Idaho, mostly collected 
by Coquillett and Aldrich. 

160 Annals Entomological Society of America [Vol. IV, 

Proctacanthus coquillettii n. sp. 

Total length 25 to 27 millimeters. Mystax and beard white or very- 
pale yellowish, palpi black with white hair, antennae dark, nearly black, 
third segment short, oval, arista nearly three times as long as the seg- 
ment which bears it. Thorax dark, brownish gray poUirtose and with 
black and gray hair; legs with black and gray bristles and gray hair; 
femora uniformly dark, nearly black, tibiae and tarsi brownish red ; wing 
hyaline with the middle third quite distinctly brown, abdomen imi- 
formly dark, gray pollinose and with numerous gray hairs. Male geni- 
talia rather short, appendages and other parts margined with dense 
rows of gray hair. Female genitalia with the end piece densely pilose 
but devoid of a tenninal circlet of enlarged spines. 

It is easily known by the brown patch on the middle of each 
wing, by the short third antennal segment and by the gen- 
italia of the male. The female genitalia agree rather closely 
with occidentalis but differs from all other species by the 
spines at the tip being small and not plainly differentiated 
from others of the last segment of the ovipositor. 

The female type and a male from Los Angeles, California, 
collected by coquillett, for whom I am pleased to name the 

Proctacanthus micans Schiner. 

Total length 26 to 3G millimeters. Mystax pale yellow, beard white, 
palpi black with most of the hairs with which they are clothed white, 
although there may be a few black ones. Occipito-orbital bristles 
mostly pale but in some specimens there are plenty of black ones. 
Thorax brown in ground color, grayish brown pollinose and with the 
usual dorsal markings, numerous black bristles on the posterior part 
with some gray hairs intermixed, anterior part with short black hairs; 
wings nearly hyaline with the veins quite distinctly margined with 
brown; legs rather dark from general view, femora dark before, brown 
behind, tibiae and tarsi in large part brown but somewhat variable. 
Abdomen dark, grayish brown pollinose; male genitalia rather small 
and compact, appendages scarcely protruding beyond the other parts, 
of nearly the same width throughout from side view and clothed with 
fine white hair. Female genitalia black or brown, end piece sparsely 
hairy and with a distinct circlet of strong spines at the tip. 

The small compact male genitalia and hyaline wings with 
brown margined veins are characteristic. 

Numerous specimens from Colorado, Arizona and New 

Proctacanthus duryi n. sp. 

Total length 26 to 28 millimeters. Mystax pale yellow, beard and 
hair of the palpi nearly white, in fact all the bristles and hairs of the 
head are white or very pale yellowish. Whole body yellowish gray 

1911] Promachus and Proctacanthus 161 

pollinose, dorsum of the mesothorax and scutellum with pale bristles; 
wing clear hyaline, veins pale brown; legs rather light colored, femora 
darker before than behind, all parts of the legs with black bristles and 
pale hair. Hair and bristles of all parts of the abdomen gray. Male 
genitalia about as long as the last two abdominal segments, female 
genitalia with a circlet of strong spines at the tip. 

The rather small size of the sjjecies, with the white bristles 
of the whole body, especially the scutellum, and the general 
pale color are distinctive. 

Male type and numerous other specimens taken along the 
Ohio River in Ohio and Kentucky by Charles Dury for 
whom the species is named. 

Proctacanthus arno Townsend. 

Total length 28 to 36 millimeters. Mystax white, usually with a few 
black bristles beneath, beard white, palpi black with black and white 
hairs, proboscis black, occipito-orbital bristles partly black and partly 
white, whole body grayish brown pollinose. Thorax with the usual 
markings above, mesothorax above with short black hairs anteriorly 
and rather stout black bristles posteriorly, scutellum with many black 
bristles and a few white hairs anteriorly. Legs in general color rather 
dark, femora nearly black before, brown behind, tibiae and tarsi some- 
what variable but often quite dark so that they show no contrast with 
the femora; wings hyaline, usually with the veins faintly margined 
with brown, but in teneral specimens they are entirely glassy clear. 
Abdomen rather slender, male genitalia elongate, longer than the last 
two abdominal segments, appendages from side view, narrowest at the 
middle, slightly widened and curved inward at the tips so that from 
dorsal view a distinct open space is enclosed beyond the other genital 
parts. Female genitalia usually shining black with short white hairs, 
end piece rather sparsely hairy with a circlet of strong black spines at 
the tip. 

The long male genitalia as described characterizes this 
species fully. 

The type was collected at the southern end of Lower Cali- 
fornia. Specimens are at hand from southern Arizona, New 
Mexico, Utah, Palo Alto, California, and El Paso, Texas. 

Proctacanthus nigrofemoratus n. sp. 

Total length of the male type 30 millimeters, other specimens vary 
from 23 to 30 millimeters. Mystax beard and hair of the palpi white; 
palpi, antennae and proboscis black; occipito-orbital bristles partly 
black and partly white. Thorax gray pollinose, mesothoracic dorsum 
with black hairs and bristles, scutellum with black bristles; wings 
hyaline with pale veins. Femora uniformly black all over, tibiae pale 
on the basal parts, otherwise dark, nearly black, as are all the tarsi. 

162 Annals Entomological Society of America [Vol. IV, 

Basal part of the abdomen largely dark in ground color and gray pol- 
linose, last three segments red in ground color and gray pollinose. 
Male genitalia slightly shorter than the last two abdominal segments 
combined, rather small and compact; appendages red, from lateral 
view of nearly the same width throughout and rounded at the apex. 

The entirely hyaline wings, the uniformly black femora and 
the small size of the specimens make the species easy of deter- 
mination. The female genitalia shining black with a distinct 
circlet of spines at the tip. 

The male type and two other males and four females 
from San Jose, Mexico, collected by D. L. Crawford and 

Proctacanthus philadelphicus Macquart. 

Total length 2S to 3(3 millimeters. Mystax dark yellow, usualh- with 
some black biistles intennixed, beard slightly paler than the mystax, 
palpi black and mostly with black hairs; occipito-orbital bristles part 
black and part yellow. Thorax rusty brown pollinose, mesothoracic 
dorsum with short black hairs anteriorly and with black bristles and 
hairs posteriorly, scutcllum with numerous black bristles and hairs and 
often with a few white hairs intennixed. Legs in most part brown 
with black bristles and pale hairs, femora darkest anteriorly, wings 
uniformly brown all over. Abdomen rusty brown pollinose. Male 
genitalia slightly shorter than the last two abdominal segments com- 
bined, appendages brown, from side view nearly straight and near the 
same width throughout, the tips rounded but not curved towards each 
other. Female genitalia shining black, end piece rather bread, sparsely 
hairy and with a circlet of strong black spines. 

The intense brown color of the body and wings and the 
dark yellow mystax are characteristic of the species. 

Specimens from New Jersey, Maryland, Connecticut, Mas- 
sachusetts, Virginia and Pennsylvania. 

Proctacanthus longus Wiedemann. 

Total length 32 to 36 millimeters. Mystax yellow, beard paler than 
the mystax, palpi black with paler hairs, occipito-orbital bristles part 
black and part pale. Thorax brown, gray pollinose, mesothoracic 
dorsum with the usual markings rather plainly shown, anterior part 
mostly with short black hairs, posterior part with black bristles, scutel- 
lum with black bristles and hairs; legs brown with black bristles and 
numerous pale hairs; tibiae slightly lighter than the femora; wings 
uniformly brown, wide and rather long. Abdomen brown, brownish 
gray pollinose, male genitalia shorter than the last two abdominal seg- 
ments combined, from side view appendages nearly straight, of nearly 
tmifonn width with the tips rounded but not curved toward each other 
when viewed from above; female genitalia black with a terminal circlet 
of strong spines. 

1911] Promachus and Proctacanthus 163 

Related to philadelphicus but the pale hairs of the palpi, 
long, wide wings and more grayish pollinosity of the body 
serve to characterize it. 

Proctacanthus craverii Bellardi reported from Mexico, 
from the description, I take to be a synonym. 

Specimens at hand from Georgia, Florida and Texas. 

Proctacanthus milbertii Macquart 

Total length 28 to 40 millimeters. Mystax pale yellow, beard paler 
than the mystax, palpi black with black hair. Thorax gray pollinose, 
mesothoracic dorsum with the usual markings rather plainly shown, 
scutellum with black bristles and usualh' with a few pale hairs; wings 
uniformly light brown; legs brown, tibiae and tarsi lighter than the 
femora which are darker anteriorly than behind. Abdomen gray or 
in some cases brownish gray pollinose. Male genitalia shorter than the 
last two abdominal segments combined, appendages not curved toward 
each other at the tips; ovipositor usually black but occasionally red or 
partly red, with a circlet of strong spines at the tip. 

Related to philadelphicus, but the mystax and wings are 
paler; the abdomen is gray and not brown pollinose and the 
thoracic markings are rather more distinct. 

Specimens from western Ohio, Indiana, Wisconsin, Kansas, 
Missouri, Michigan, Colorado, California and British Colum- 
bia. It is one of our most common and widely distributed 

Under the name Asilus missouriensis, Riley has written of 
this species as a distinct enemy of bees. 

Proctacanthus nigriventris Macquart. 

Total length 30 to 35 millimeters. Body very dark, nearly black; 
wings unifomily rather pale brown. Mystax very pale yellowish, often 
with black bristles intermixed, beard gray, occipito-orbital bristles 
black. Thorax dark, brown pollinose, mesothoracic dorsum with 
black hair and bristles, usual markings not plainly differentiated; scu- 
tellum with black hairs and bristles. Wings uniformly rather pale 
brown, legs dark nearly black; femora, except the apices, black, apexes 
of femora, tibiae and tarsi dark brown . Abdomen dark, nearly black, dark 
pollinose with a narrow band before each incisure light pollinose, venter 
and sides of the last two or three segments in the male gray pollinose; 
male genitaHa dark red, rather short, appendages widest at the base, 
gradually narrowed toward apex where they are evenly rounded; 
ovipositor shining black with strong black spines at the tip. • 

The very dark color of the whole body is distinctive. 
Specimens from New Jersey, collected by H. S. Harbeck, of 

164 Annals Entomological Society of America [Vol. IV, 

Proctacanthus brevipennis Wiedemann. 

Total length 20 to 28 millimeters. Mysta.x usually white but 
sometimes yellowish and occasionally largeh' black, in any case there 
is likely to be more or less black bristles mixed with the others, beard 
white, palpi with black hair, occipito-orbital bristles mostly black. 
Thorax brown, gray pollinose, mesothoracic dorsum with the usual 
markings very plainly differentiated, anterior part with short black 
hairs, posterior part with numerous black bristles; scutellum with black 
bristles and a few white hairs before. Wings tmiformly pale brownish; 
legs brown, tibia and tarsi lighter than the femora, which are nearly 
black before and brown behind. Abdomen rather dark colored, more 
or less gray pollinose, male genitalia red, rather short, small, of nearly 
the same width throughout and evenly rounded at the tips. 0^'i- 
positor shining black, except occasionally when it is more or less red, 
with a distinct circlet of spines at the tip. 

The small size and plainly differentiated thoracic markings 
are usually sufficient to place specimens of this species. 

Specimens at hand ffom North Carolina, Kansas, New 
Jersey, Florida, and Georgia. The type of the species was 
taken in Kentucky. 


In 1838 Macquart described the genus Trupanea to receive 
the species which fall here. Ten years later Loew discovered 
that Macquart's name could not stand on account of previous 
usage and proposed Promachus to take its place. The species 
are large, although somewhat smaller than the species of Proc- 
tacanthus. The bristles of the antennae are naked, the veins clos- 
ing the discal and fourth posterior cells are not parallel, the pos- 
terior branch of the third vein terminates beyond the tip of the 
wing, there are three submarginal cells and the abdomen is 
longer than the wings. These characters will place the species 
easily in most cases but certain species of Mallophora may cause 
trouble. In case they do the fact that the members of the lat- 
ter genus have obtuse claws, while those falling under Pro- 
machus have pointed claws should obviate the difficulty. 
There are at least nineteen valid species of the genus described 
from North America, but since five of these from Mexico and 
Central America could not be procured only fourteen are 
treated in this paper. The genus appears to reach its greatest 
development to the southward and I have reasons for believing 
that there are yet in North America several undescribed 

1911] Promachus and Proctacanthus 165 


1. First subniarginal cell with a gray shadow which sometimes is very narrow. .2 
First submarginal cell without any trace of a gray shadow 12 

2. Male genitalia with dense silvery hair above, color of the tibiae very near 

like that of the femora 6 

Male genitalia without silvery hair above, color of the tibiae usually in strik- 
ing contrast to that of the femora 3 

3. Gray bands on the dorsum of the abdomen fully as wide as the black bands. .4 
Gray bands on the dorsum of the abdomen confined to narrow posterior 

borders of the segments 5 

4. Thorax deep reddish brown, male genitalia clothed with black hair.ruflpes 
Thorax yellowish gray, male genitalia clothed with gray hair. . . .vertebratus 

5. Gray shadow in the first submarginal cell less than half as wide as the cell; 

male genitalia rounded at apex sackeni 

Gray shadow in the first submarginal cell darker in color than usual, distinct- 
ly more than half as wide as the cell; male genitalia truncate at apex forfex 

6. Abdomen with black hair on some or all of the first five segments above. .9 
Adbomen without black hair on any of the segments above 7 

7. Abdomen uniformly clothed with short gray hair, mystax very pale yellow or 

nearly white, beard white. truquii 

Abdomen clothed with yellow hair, beard and mystax distinctly yellow. . . .8 

8. Gray shadow in the first submarginal cell wider than the marginal cell, male 

genitalia almost as long as abdominal segments five, six and seven com- 
bined fitchii 

Gray shadow in the first submarginal cell distinctly narrower than the margi- 
nal cell, male genitalia shorter than abdominal segments six and seven 
combined quadratus 

9. Mystax white or with a very slight shade of yellowish, legs chiefly black . . 10 
Mystax plainly yellow, legs chiefly yellowish red 11 

10. Abdomen densely brown pollinose on the sides albifacles 

Abdomen gray pollinose on the sides and extending across narrowly on the 

hind margins of the segments princeps 

11. Abdomen densely brown pollinose on the venter and sides, wings plainly 

brownish bastardii 

Abdomen pale yellowish gray pollinose on the venter and sides, wings nearly 
hyaline quadratus 

12. Legs pure black , 13 

Legs more or less reddish 14 

13. Abdomen distinctly banded with black and gray, the gray bands slightly 

more than half as wide as the black nigripes 

Abdomen not distinctly banded, although the narrow posterior margins of 
the segments are somewhat different from the other parts by appearing 
whiter aldrichil 

14. Very large species, 35 to 40 millimeters in length giganteus 

Small species, not over 25 millimeters in length minusculus 

Promachus vertebratus Say. 

Total length 26 to 31 millimeters. Front and face clothed with 
yellow pollen, hairs and bristles of the whole head yellowish, beard 
paler than the mystax, often some of the bristles on various parts of the 
head are black but there does not appear to be any uniformity as to 
which ones are black and often there are none of this color. Proboscis 
shining black, palpi clothed with yellowish hair with an intermixture 
of black in some specimens, antennae black. Thorax yellowish pol- 
linose, dorstmi with a brown stripe at the middle divided by a narrow 
gray line, wings dilute brownish; legs somewhat variable in color, 
clothed with black bristles and recumbent pale hair, femora in large 
part black although they may be almost entirely reddish, or even only 

166 Annals Entomological Society of America [Vol. IV, 

reddish in part ; tibiae reddish with the extreme apices darker ; all of the 
tarsi brown or black. Abdomen largely gray pollinose and clothed with 
pale hairs; dorsally the first six segments each have a rectangular black 
marking which occupies about the anterior half of its respective seg- 
ment. Male genitalia black, clothed with pale hair, female genitalia 
shining black. 

, Specimens from Kansas, Wisconsin, Illinois and Ohio. 

Promachus rufipes Fabricius. 

Total length 28 to 35 millimeters. This is a larger species than 
vertebratus. The color of the thorax is much browner than in that 
species and the color of the wings is more pronounced. The palpi are 
black with black hair and the hypopygium is shining black with scat- 
tering black hairs. The legs are somewhat variable, or at least in the 
specimens before me they show two distinct types of coloration. The 
specimens that agree closer to the original description have pure black 
femora and reddish yellow tibiae with black apices. In another set of 
specimens the femora and tibiae, except the apices, are reddish brown 
throughout and do not show any contrast. In other respects the two 
series of specimens exactly agree and I prefer to consider them all 

Specimens at hand from District of Columbia, Tennessee, 
Mississippi, Iowa and Ohio. 

Promachus sackeni n. sp. 

I take this to be Promachtis No. 2, Biolog. Cent. Am., 
Volume I, page 193. 

Total length 22 to 27 millimeters. Mystax composed of black and 
yellow bristles intermixed, palpi black, mostly with black hairs but 
there may be light colored ones intermixed, especially on the basal 
parts, beard white, thorax gray pollinose above with the usual mark- 
ings, wings uniformly pale brownish, the dark marking in the first 
submarginal cell very little more than one-fourth as wide as the cell at 
the widest part, femora, apex of the tibiae and the tarsi black with 
black bristles, remainder of the tibiae yellowish red with numerous 
pale hairs and a few black bristles, claws pointed and distinctly curved. 
Abdomen with much yellowish hair which on the sides is rather long; 
segments two to seven in the male and two to five in the female ■with 
gray pollinose triangles on the sides, the inner angles on each side 
ineeting above on each of segments two to five in both sexes and fonning 
a posterior gray band; the hypotenuse of each triangle is slightly con- 
cave, leaviiag on each segment a semicircular spot which is clear black; 
hypopygium black, slightly wider than the abdomen, evenly rounded 
posteriorly, with mostly black hairs dorsally and pale yellow hairs 
laterally; ovipositor black, somewhat longer than abdominal segments 
four and five. 

1911] Promachus and Proctacanthus 167 

Three males and three females from southern Arizona, in 
July and August, some of them collected by the late Dr. F. H. 

Osten Sacken's specimens were procured in northern Sonora, 
Mexico, by Morrison. In a note after his description he men- 
tioned Promachus trapezoidalis Bellardi as closely related, but 
Williston has shown, rightly I think, that the latter belongs to 
Mallophora. I have seen a male of this from Brownsville, 
Texas. Bellardi mentions the blunt claws and Williston calls 
attention to the same character. 

Promachus forfex Osten Sacken. 

This is a new name for Promachus quadratus Bellardi, 
since there is an older quadratus by Wiedemann. 

Total length 23 to 30 millimeters. Mystax largely composed of 
yellow hair, but a few of the finer ones are black, antennae and pro- 
boscis black, palpi with black hair, occipito-orbital bristles black. 
Thorax sparsely gray pollinosc with the usual darker markings dorsally, 
scutellum with two more or less irregular rows of black bristles and 
numerous shorter yellow hairs; wings uniformly yellowish brown, dis- 
tinctly darker than in sackeni, cloud in the first submarginal cell two- 
thirds as wide as the cell, dark colored and with a distinct outline; 
femora, tips of the tibiae and tarsi, including the claws pure black, 
remainder of tibiae and pulvilli yellow, balancers pale 3'ellow; abdomen 
largely clothed with light yellow hair, sides and narrow posterior mar- 
gins of the first five segments in the female and seven in the male gray 
pollinose leaving a large quadrate dark marking abo\'c on each of the 
segments. Ovipositor composed of three segments, shining black; 
hypopygium black with black hair, from dorsal view gradually widened 
toward the apex where it is plainly truncate. 

Male and female from Cordoba and a female from Vera- 
cruz in Mexico, collected by D. L. Crawford. Also a female 
from Puerto Barrios, Guatemala, collected by E. B. Williamson. 
The species appears to be southern in distribution and is not 
likely to be taken in the United States. 

Promachus fitchii Osten Sacken. 

Fitch described this species under the name apivora because 
he found it injurious to bees. His name could not stand, how- 
ever, as it had been used before in the genus. 

Total length 25 to 30 millimeters. Mystax, beard and hairs of the 
palpi distinctly yellow, occipito-orbital bristles pale yellow intermixed 
with black ones which are more numerous in some specimens than in 
others. Thorax yellowish brown pollinose above with short black 

168 Annals Entomological Society of America [Vol. IV, 

hairs anteriorly and longer black hairs and bristles posteriorly, a tuft of 
yellowish hairs before the scutellum in well preserved specimens, scu- 
tellum with numerous yellow hairs and a few black bristles ; legs mostly 
yellowish red although usually there is a black or blackish area on the 
front side of each femur; hairs of the legs yellow, bristles black; wings 
with a pale brownish tinge, shadow in the first submarginal cell plainly 
evident, a little more than a third of the width of the cell at the widest 
part, ground color of the abdomen black, unifonnly clothed with rather 
short yellow hairs all over. 

The long hypopygium is clearly distinctive in this species. 
It is nearly twice as long as in any other species I have seen. 

Numerous specimens from Clay County, Kansas, collected 
by Prof. J. H. Schaffner. Aldrich catalogues it from Nebraska, 
Missouri, Kansas, Connecticut and Florida. 

Promachus albifacies Williston. 

Total length 22 to 28 millimeters. Mystax and beard white, at 
most only a few black hairs on the oral margin; palpi with many black 
hairs and usually with some scattering white ones among them; rear of 
the head with white hairs and a few black bristles above. Thorax 
brown pollinose with the usual stripes above, a tuft of white pile in 
front of the scutellum largely surrounded by black bristles, scutellum 
with numerous white hairs and black bristles; wings nearly hyaline, 
gray shadow in the first submarginal cell less than one-third as wide as 
the cell at the widest point; legs largely black but somewhat variable, 
often the femora are red posteriorly and the same may be said of each 
tibia at base. Abdomen pure black in ground color, distinctly brown 
pollinose on the venter and sides and on each side of each segment are 
numerous white hairs which tend to reach across the dorsum and form a 
very narrow hind border to the segment. Hypopygium small, short 
and clothed above with abundance of silver white pile. 

The species is known from California, Colorado, Arizona, 
New Mexico and Mexico. 

Promachus princeps Williston. 

Total length 27 to 32 millimeters. Mystax white with a few black 
bristly hairs on the oral margin, beard white, palpi with numerous 
black hairs and a few white ones, rear of the head with white hair and 
some black bristles above; thorax gray pollinose, wings hyaline, gray 
shadow in the first submarginal cell very narrow and usually not sharply 
defined, legs approaching black, each femur posteriorly and each tibia 
at the base more or less dark reddish; abdomen black in ground color, 
gray pollinose on the sides and venter, with an area of white hairs on 
each side of each segment and these tend to approach each other and 
unite and thus fdnn a band across the posterior margin of the segment. 
The abdomen is somewhat variable when different specimens are 
compared on account of the area covered with white hair. 

1911] Promachiis and Proctacanthiis 169 

This species may be separated from albifacies by the gray 
instead of brown pollinosity of the thorax, and sides and venter 
of the abdomen, by the wings being more nearly hyaHne and 
by the gray shadow in the first submarginal cell being narrower 
and less sharply defined than in that species. 

The species is known from CaHfornia, Oregon and Wash- 

Promachus bastardii Macquart. 

Asilus laevinus, Trupinea rubiginis and Asilus ultimus of 
Walker, and Promachus philadelphicus Schiner are considered 

Total length 21 to 2S millimeters. Mystax and beard yellow, the 
latter paler than the fonner, palpi clothed with black hair, occipito- 
orbital bristles black; thorax brown, the usual markings present on the 
dorsum but not very plainly shown, legs in general color red with white 
hair and black bristles; in most specimens there is a black stripe of 
greater or less width on the anterior side of each femur, and the front 
legs especially may have more or less yellow pile on the tibiae and 
tarsi; wings quite distinctly unifonnly brownish, shadow in the first 
submarginal cell about half as wide as the cell at the widest part, not 
very dark gray but with definite limits. Abdomen distinctly brown 
on the venter and sides and clear black above. From dorsal view the 
abdomen shows a very narrow band of white hair on the posterior 
margin of each segment and black hairs on a clear black background 

The hypopygium is clothed with silvery white hair above 
but is distinctly wider than in albifacies and princeps. 

Specimens have been taken in New York, Massachusetts, 
Pennsylvania, Michigan, Kansas, Georgia and Ohio. 

Promachus quadratus Wiedemann. 

Total length 22 to 29 millimeters. Mystax and beard yellow, the- 
latter distinctly lighter than the former, palpi black with yellow hair, 
but with a few black ones intermixed, rear of the head with yellowish 
hair, occipito-orbital bristles black. Thorax light brown pollinose with 
the usual middorsal stripe and lateral markings; wings very pale yel- 
lowish all over, shadow in the first submarginal cell narrow, hardly 
one-fourth as wide as the cell at the widest place ; legs brown with light 
hair and black bristles, anterior part of each femur darker than the 
other parts of the leg; balancers brown. From above each abdominal 
segment has a rather wide posterior margin which is clothed with pale 
yellowish or white hair and which widens distinctly towards each side. 
The ground color of the abdomen is black and the color shows plainly 
on the anterior part of each segment but not so extensively as in bas- 
tardii. In the male the genitalia is of medium size and clothed above 
with silver white hair as in related species. 

170 Annals Entomological Society of America [Vol. IV, 

There has been some uncertainty in regard to quadratus but 
these specimens agree well with the original description. In 
comparison with bastardii the color as a whole is lighter, the 
palpi are clothed with mostly yellow hair instead of mostly 
black hair, the wings are much nearer hyaline, the gray shadow 
in the first submarginal cell is much narrower and the abdomen 
has much more light hair. Weidemann's type is recorded as 
from Georgia. 

Specimens are at hand from Cameron Parish, Louisiana. 

Promachus truquii Bcllardi. 

Total length 24 to 3(i millimeters. Mystax pale yellowish, beard 
white, rear of the head with white hair, occipito-orbital bristles mostly 
black, palpi with many black hairs but beneath there are a number of 
pale yellow ones; thorax brown pollinose, sides with white hairs, dorsum 
with many short black hairs anteriorly and rather long white hair and 
black bristles posteriorly ; legs in large part reddish brown with white hair 
and black bristles, anterior side of each femur largely black; wings nearly 
unifomih- pale browTiish, hardly as dark as in bastardii, gray shadow in 
the first submarginal cell plainly evident and with distinct limits, about 
a third as wide as the cell at the widest j^lace. Abdomen brown pol- 
linose on the venter and sides, black dorsally, unifonuly clothed all 
over with short white hair, no black hair except a few black bristles on 
either side of the first segment. The hypopygium is furnished with 
silvery hair above very much as in albifacies. 

The most apparent difference bewteen this and related 
species is the uniform distribution of rather short white hair 
over the abdomen and the absence of black hair except the 
black bristles on the sides of the first segment. 

Several specimens from the Huachuca Mountains, Arizona. 
Besides the species has been reported from several places in 

Promachus nigripes n. sp. 

Total length 28 to 32 millimeters. Mystax and beard white, rear 
of the eyes with white hair, occipito-orbital bristles largely black, palpi 
black with numerous black hairs and a few white ones on the lower 
side, antennae black. Thorax gray pollinose, mesothoracic dorsum 
with the usual markings not conspicuous, clothed largely with black 
hair and bristles, sides of the thorax with long gray hairs, scutcllum 
with black hair and bristles; legs black, mainly with black hair and 
bristles, but there may be some white hairs especially on the femora; 
wings hyaline, no gray shadow in the first submarginal cell. Abdomen 
from above banded with black and white, a white pollinose band clothed 
with white hair on the posterior part of each seginent and a wider 
black hand clothed with black hair before it, venter and sides white 

1911] Promachiis and Proctaciuithns 171 

I^ollinosc with white hairs. Hypopygium rather wide and clothed 
above with silvery hair, posterior ventral margin of the eighth segment 
with black and white hairs. 

There is some variation among the different specimens. 
The hair on the posterior part of the mesothorax may be largely 
white, the legs may vary in the extent of the white or black 
hair present, although the bristles are always black, and in the 
male the margin of the eighth ventral segment may be furnished 
entirely with black hairs. 

The distinct limitation of the black and white areas on the 
dorsum of the abdomen so as to form bands, together with the 
entirely black legs characterize the species. 

The male t3'pe and seven other males and females taken at 
Albuqtierqtie, New Mexico, by J. R. Watson. 

Promachus aldrichii n. sp. 

Total length 2S to 30 millimeters. Mystax and beard white, occipito- 
orbital bristles mostly black, palpi black with white hairs, only a few 
black ones above. Dorsum of the mesothorax mainly with black 
hairs, and bristles, but there is some rather long white hair on the scu- 
tellum and before it, prothorax gray pollinose, usual markings present 
but they are not very plainly shown; legs black, clothed with black and 
white hair and black bristles; wings clear hyaline, no shadow in the first 
submarginal cell ; abdomen black in ground color, gray pollinose on the 
sides and venter and clothed everywhere with rather short hair which 
is almost all white, there being only a very few short black hairs on the 
anterior dorsal part of each segment. In the male the posterior ventral 
margin of the eighth adbominal segment is furnished with a fringe of 
white hairs and the hypopygium is silvery pilose above. 

The most characteristic thing about the species is the uni- 
form distribution of the white hairs on the abdomen so that 
this part does not appear evenly banded as in nigripes. 

The male type and one female from Utah, and received 
from J. M. Aldrich of Moscow, Idaho. 

Promachus minusculus n. sp 

Total length 24 to 20 millimeters. Mystax and beard pure white, 
palpi black with some black and some white hairs, occipito-orbital 
bristles largely pale although there are always a few black ones inter- 
mixed. Thorax gray pollinose, thoracic dorsum with the markings not 
very distinct, clothed with black and white hair and black bristles, 
bristles of the scutellum partly white and partly black, wings pure 
hyaline, veins brown. Legs dark reddish in general coloration, femora 
darkest anteriorly, tibiae red, tarsi nearly black, all the parts clothed 
with reclining white hairs and black bristles. Abdomen rather dull 
black in general coloration, venter, sides and posterior part of the dor- 

172 Annals Entomological Society of America [Vol. IV, 

sum of each segment clothed with white hairs, anterior part of the 
dorsum for more than half the length of each segment clothed with 
black hairs. Hypopygiiom small and clothed both above and below 
with rather long silvery white hair, ovipositor black. 

Easily known from other species of its group by its small 
size and general rather light color as well as by the very small 
hypopygium which is densely clothed with silvery hairs below 
as well as above. 

Male type and five other males and females taken near 
Albuquerque, New Mexico, bj' J. R. Watson. 

Promachus giganteus n. sp. 

Total length, male 37, female 41, millimeters. A/fystax and beard 
white, hair of the rear of the head white, occipito-orbital bristles mostlj' 
black, palpi black, largely with white hairs, but there are some black 
ones above, antennae black; general color of the thorax reddish brown, 
mesothoracic dorsum with a rather wide middorsal black stripe which 
is divided lengthwise by a narrow red interval, on either side is a second 
black area which is divided by a narrow red space which follows the 
transverse suture. As the surface of the dorsum of the meso thorax is 
somewhat denuded in the specimens studied it is likely that the mark- 
ings described are plainer than would be the case otherwise, scutellum 
clothed with white hair and with two rows of black bristles near the 
margin; legs in large part dull reddish and clothed with black bristles 
and recimibent white hair, tarsi nearly black, femora darkened anteri- 
orly; wings hyaline, no gray shadow in the first submarginal cell, veins 
brown and some of them towards the apex very narrowly margined 
with a nearly obsolete brownish shade. Abdomen as seen from above 
alternately banded with black and white, the former color usually 
wider than the latter. The black and white is segmentally arranged, 
the anterior part of each segment is black with black hair and the pos- 
terior part is white pollinose with white hair, sides and venter of the 
abdomen corresponds in color with the posterior parts of the seg- 
ments. In the male the hypopygium is silvery pilose above and the 
posterior ventral margin of the eighth abdominal segment is furnished 
with a thickly placed row of white hairs. 

The species cannot be mistaken among described North 
American species on account of its very large size. 

The type female and one male taken at El Paso, Texas, by 
D. L. Crawford, and received from Carl F. Baker, of Pomona 
•College, Claremont, California. 



By A. F. Burgess. 
Bureau of Entomology, Melrose Highlands, Mass. 

Among the interesting data in the report on "The Gypsy 
Moth," by Forbush and Fernald, which was published by the 
Massachusetts State Board of Agriculture in 1896, will be found 
a record of a series of experiments to determine the distance 
which young gypsy moth caterpillars will travel. Seven cater- 
pillars were used, and it was found that the distance they 
crawled before dying ranged from 36 to 144 feet. Another 
record of experiments conducted with another insect is given 
on page 23 of bulletin 72 of the New York State Museum, which 
is bulletin 19 on entomology, published in 1903. In it Dr. 
Felt writes concerning the grape root worm that he carried on 
"some experiments to determine the burrowing and traveling 
power of these little creatures. One small grub was placed on 
a piece of paper at 9.27 in the morning and its wanderings were 
carefully traced with a pencil till 4.43 in the afternoon. The 
little creature traveled almost continuously during the entire 
period and showed a decided tendency to turn to the left. It 
covered the relatively enormous distance of over 47 feet in 
seven hours, or an average of about 2 yards an hour. The 
grub was placed in a dry vial and under such unfavorable 
conditions lived about three days." 

These experiments show that insect larvae are able to sur- 
vive for a much longer time than would be expected, and to do 
so under adverse conditions. The ability to travel until food is 
secured being absolutely necessary to the existence of the indi- 
vidual this factor plays a very important part in determining 
the ability of larvae to survive, and especially is this true of 
predaceous forms. Our attention was strongly directed to this 
matter in connection with the work of importing and colonizing 
certain predaceous enemies of the gypsy and brown-tail moths 
in Massachusetts. The ability of the larvae of Calosoma 
sycophanta, a predatory species which has been received from 
Europe, to travel any great distance in search of food is of para- 

_ • Read at the Minneapolis meeting, Dec. 28, 1910, in joint session with the Association of Economic 



174 Annals Entomological Society of America [Vol. IV, 

mount importance to the well-being of the species, and al- 
though the insects had been found to be very active in breeding 
jars it seemed worth while to secure more accurate data as to 
their powers of locomotion. Accordingly, a set of experiments 
was planned to determine the distance that a larva of this 
species would travel from the time of hatching until it died, 
provided no food or moisture was supplied. In order to carry 
out the test it was necessary to do so under conditions which 
never occur in nature, but the results indicate the practical 
impossibility of carrying on tests over so long a period unless 
absolute control is secured, so that a careful record can be made. 
The following apparatus was used (Plate XIII) : A small 
table 3 ft., 8 in. long, by 2 ft. wide was provided with spools 
at each end near the top, so that a roll of paper could be reeled 
across the top of the table, the result being accomplished by 
turning the spools. Beneath this paper was placed a piece of 
stiff wrapping paper which extended beyond the sides of the 
paper connected with the reels, and the edges were bent upward 
in such a manner as to prevent the escape of the larva from the 
sides of the tabic. The paper on the reels was ordinary wrap- 

Fig. I. Rolls of Records. 

Annals E. S. A. 

Vol. IV, Plate XIII. 

A. F. Buvfiess. 


Larva of Calosoma Sycophanta. 


ping paper 18 inches wide. The larva was placed in the center 
of the table and a record of its travels was made with a lead 

At 8:30 A. M., June IS, a newly hatched larva was placed in 
the center of the table bj^ Mr. C. W. Collins, and the record was 
kept throughout the day with the assistance of Mr. R. G. 
Smith. The table had been placed in an attic room in the lab- 
oratory, having only one window, which was on the west side 
of the house. During the day the larva moved rapidly away 
from the light, and when it had reached the end of the roU the 
table was turned end for end and the insect began traveling 
in the opposite direction. Although there were forty yards 
of paper on this roll, it was necessary at 1 p. M. to remove the 
larva and substitute a fresh roll, as it had crossed the paper 
from end to end five or six times. 

The complete record of the travel of this larva required 11 
rolls of paper and one assistant and sometimes two, depend- 
ing on the activity of the insect, had to be constantly at work. 
(Fig. 1.) The experiment was carried through continuously 
until the larva died, and the extraordinary vitality which it 
exhibited, promised, at one time, to exhaust the supply of assist- 
ants that could be spared for the work. 

The following table gives the data secured from the record 
on each roll. 




Rate per min. 

8 30 a. m to 1 p m. 

i\i hrs. 
3 2-3 hrs. 
5 hrs. 

1323.63 ft. 

719.40 ft. 
1213.14 ft. 
1164.51 ft. 

733.17 ft. 

926.03 ft. 

933.15 ft. 

712.15 ft. 

575.91 ft. 
766.5 ft. 

190.44 ft. 

4 9 ft 

1 p. m. to 4.40 p. m 

4.40 p. m. to 9. 40 p. m 

3.27 ft. 
4.04 ft. 

9.40 p. m. to 1.45 a. m 

4' 2 hrs. 

6 hrs. 

3 hrs. 
4 23-30 hrs. 

5M hrs. 
19^4 hrs. 

4.75 ft. 

2.03 ft. 

S.40 a. m. to 11.40 a. m. 

4.03 ft. 

11.54 a. .m to 4.40 p. m 

5 p. m. to 10.15 p. m , 

3.26 ft. 
2,26 ft. 

10.35 p. m. to 2.05 a. m 

2.15 a. m. to 1.30 p. m 

2.74 ft. 
1.13 ft. 

1.30 p. m. to S.45 a. m 

.16 ft. 

The larva remained alive from 8:30 Saturday morning 
until 8:45 Tuesday morning, 72 hours, and was active the greater 
part of the time. It was necessary to place the insect in a glass 
each time a new roll of paper was attached, so that the entire 
length of time that the larva was actually on the paper was about 
70 hours. The table gives the distance traveled during differ- 
ent periods, the total amounting the 9,058 feet, or 1.71 miles. 

176 Annals Entomological Society of America [Vol. IV, 

The highest rate of travel per minute was during the first 4}/^ 

hours, and averaged 4.9 ft. For the first 24 hours the average 

was 3.69 ft. per minute; during the next S hours the average 

dropped slightly and for the remaining period the average was 

gradually reduced until the larva died. 

The temperature in the room where the experiment was 

conducted ranged considerably higher than that outside the 

building, the following records being taken from the report 

of the U. S. Weather Bureau at Boston. 

— June 18 — Maximum, 79 degrees; Average, 70 degrees. 

" 19— " 75 " " 68 

" 20— " 86 " " 74 

" 21— " 92 " " 80 

The larva was kept continually on a dry surface, so that 
evaporation was very rapid, and doubtless reduced the length 
of time that the insect would have survived under normal con- 

For the first 36 hours the larva traveled almost continuously, 
stopping only occasionally for a minute or so to rest. 

During this time it traveled away from the light, viz., 
from the window during the day time, and at night away from 
the single electric light which was in the room. On the second 
night, which was after 36 hours of continuous travel the larva 
began traveling toward the light, and continued to do so during 
the night. 

Mr. R. G. Smith, who was on duty at the time, noted that 
from 8 A. M. on June 20, the larva traveled at a continuously 
decreasing speed. Marks were made showing direction of 
travel, and every fifteen minutes the time was noted on the 
lines of travel. Resting periods: 4.32 p. M., 8 minutes; 4.45 
p. M., 5 minutes; 5.15 P. m., 10 minutes; 5.46 P. M., 12 minutes; 
6.36 P. M., 4 minutes; 6.42 p. m., 7 minutes. The larva moved 
only several inches at a time. In the morning the larva rested 
often, but only for a few minutes at a time. Mr. J. J. Culver 
noted that about 11 p. m., June 20, the larva began to crawl 
backwards, at intervals bending the head down as if trying to 
bite itself. It would do this for two or three minutes, then either 
rest or crawl in the normal manner. At this time the larva 
had become either insensible to light, or too weak to contin- 
ually travel from it, as it traveled toward the light as much, 
if not more, than from it. In backing the larva always moved 
from the light. 


Larva of Calosoma Sycophanta. 


At 12.45 A. M., the larva was very weak, it had scarcely 
moved during the last hour, and remained in a humped-up 
position. If touched with a pencil point, it would jump, but 
would not move either forward or backward. Between 8.25 
p. M., June 20, and 1.40 A. M., June 21, the larva rested 4 hours, 
15 minutes. 

TtTT — n-Ttrml-l l ll I I i r-t n t n. ' 

Fig. 2. Distance traveled by larva during the last twenty-four hours. Fig- 
ures at left indicate number of feet each small horizontal space indicates a fifteen 
minute interval. 

Mr. H. E. Smith, who was on duty at the time, noted that 
the larva did not move from 1.45 a. m. to 2.30 A. M., when it 
backed J^ of an inch, remaining thus until 3.25 A. M., when it 
turned half way round in its tracks. The line from 1.45 A. M. 
to 4 A. M. was all backed over, except in a small circle which was 
traveled in the normal , manner. At 3.55 A. m. the larva fell 

178 Annals Entomological Society of America [Vol. IV, 

on its back and remained so until 4.05 A. M. when upon being 
touched in order to test life it grappled the lead pencil point 
and again regained its footing. It moved and backed in a 
small circle at 5.15 A. M. At 7.05 A. M. it fell on its back again, 
and remained there until it died at 8.40 A. m., June 21. 

The rate of travel from 8.30 A. M., June 20, to 8.40 A. M., 
the following morning, which was the last twenty-four hours 
of the insect's life, is shown in the diagram (Fig. 2.) and 
illustrates how remarkable was the activity until almost the 
close of its life. ■ 

No delicate balances were available for weighing the larva 
after it hatched, but as soon as it died it was weighed by Mr. 
W. G. Fall, of the Massachusetts Department of Weights and 
Measures in Boston. The average weight of ten newly hatched 
larvae was .2973 grains; the weight of the dead larva was .18 
grains, which indicates that it lost .1173 grains during the 

Although the distance traveled is probably much greater 
than what would actually occur in nature, it is remarkable that 
so much latent energy can be stored up in an egg of one of these 
beetles. It is interesting to compare the ability to travel 
possessed by this larva with that of some of the higher animals. 
Through the courtesy of Dr. W. E. Castle and his assistant 
Mr. Detlefson, of the Bussey Institution, of Harvard Univer- 
sity, we have been able to secure measurements and weights of 
young rats and guinea pigs and this information is used in mak- 
ing the comparison. The average length of all the legs of the 
larva of Calosoma sycophanta, ten specimens having been meas- 
ured, was 3.37 mm., that of a young rat, one to three days old 
twelve specimens having been measured, was 19.08 mm., or 
5 2-3 times greater, while similar measurements of a guinea pig 
showed that the average length of leg was 76.25 mm., or 22,^2 
times greater than that of the beetle larva. If the ability to travel 
of the higher animals mentioned equalled that of sycophanta, 
the rat would be able to cover nine miles without food or water, 
while the guinea pig would have to cover over 38 miles under 
the same conditions. It might be said that the locomotive 
powers of a young guinea pig are superior to that of a rat of the 
same age but doubtless either would die before traveling a very 
short distance to obtain food. 

[1911] Larva of Calosoma Sycophanta. 179 

If the traveling capacity is compared on the basis of weight 
the difference is much greater. The average weight of the beetle 
larva is 19.8 m. g., that of the rat 5250 m. g., and that of the 
guinea pig 7500 m. g. Thus if the rate of travel was in the 
same proportion the rat would cover 453 miles and the guinea 
pig 648 miles while the larva traveled 1.71 miles. 

The length of life of the insect is also worthy of note when it is 
remembered that no food or water was supplied, as it indicates 
that the young larvae are able to survive several days in the 
field without food and still have sufficient vitality to make a 
thorough and active search for their prey. 


Mr. J. B. Smith: The thing that interested me most is the 
appropriateness of the German name, "laufkafer. " Mr. Bur- 
gess' paper shows that the German name is absolutely correct 
when applied to the larva of this insect, because it certainly 
can run. 

Mr. Howard: It is estimated that there are now over a 
million of these insects in Massachusetts, that next year there 
will be ten millions, and the year after one hundred millions. 

Mr. J. B. Smith: Dr. Howard's remark recalls to my mind 
the following experience: In south New Jersey there was one 
season an outbreak of a species of Geometrid caterpillar, which 
covered the scrub oaks in enormous numbers, and that season, 
in company with two or three entomologists, I was down in the 
region where the insects occurred. We found Calosoma of 
two species, willcoxi and scrutator, were present in enormous 
numbers on the young oak trees, feeding upon these larvae. 
The next year you could look through the whole territory and 
could not find a single specimen of either species. 

Mr. Burgess: I think a possible explanation might be 
that, as far as we know the larvae of Calosoma willcoxi and 
scrutator do not cHmb trees, while the larva of sycophanta 
does so very readily. 

Mr. Summers: This certainly is an interesting paper, but 
the point that is perhaps the most interesting is the comparison 
of the rat and the guinea pig. It seems to me that is a fallacy 

180 Annals Entomological Society of America [Vol. IV, 

that ought to be pointed out. If a man could jump, in compari- 
son to his size, as far as a flea, he could jump over a church 
tower. One might say too much on the relative endurance of 
animals, of the distance they can travel, unless the animals 
are of the same size, or unless the law of reduced power of 
locomotion with added weight and size is taken into account. 
The rat with the same endurance, I believe, could not travel 
the distance stated in the paper. 

Eriocampoides limacina Retz. 

By R. L. Webster. 

Recent Injury. Nearly every year cherry and plum trees 
in Iowa suffer much injury by the common pear-slug. While 
the control of this insect has never been considered a difficult 
problem, yet it happens frequently that foliage is greatly dam- 
aged before one is aware that any slugs are present. 

Serious damage is sometimes caused to cherry trees. Young 
cherry trees in the town of Ames have been killed as a result of 
defoliation following the injury bj^ the slugs. 

Appearance of the Injury. The slugs feed on the upper 
surface of the leaves, eating the parencyhma and leaving only 
the veins and the lower epidermis. Leaves thus eaten, dry, 
turn brown, curl up and fall from the tree. Frequently in- 
fested trees are left entirely bare of foliage in midsummer. 

The Insect's Appearance. The insects in question are 
dark, olive colored slugs, with a slimy covering. The fore part 
of the body, just back of the head, is broad, but it tapers back 
of this. Where the slugs are common a peculiar sour odor 
may be noticed. 

The slugs molt five times, and when mature they are about 
2-5 of an inch long. After the last molt the slug has a clean and 
dry skin, quite free of slime, and orange in color. These orange 
slugs go to the ground and form small earthen cells in which to 
pupate. When the adult sawflies emerge they deposit eggs in 
the tissue of plants on which the young slugs are to feed. 

past history and distribution. 

This insect has long been known in Europe. As far back as 
1740 Reaumur gave an account of the pest and its injury. The 
first American account was written by Prof. Wm. D. Peck, of 
Harvard, and published in 1799. 

Professor Peck wrote an essay on the insect, entitled the 
"Natural History of the Slug- Worm, " which won for him a 
gold medal and a prize of fifty dollars, given by the Massachu- 

* Read at the Minneapolis Meeting, Dec. 28. 1910, in joint session with the Association of Economic 


182 Annals Entomological Society of America [Vol. IV, 

setts Society for Promoting Agriculture. The essay was 
printed by the Society but it has now become very scarce. 
Harris, writing in 1841, said that the pamphlet was then "out 
of print and rarely to be met with." It was the good fortune 
of the writer to find this paper of fourteen pages in the Boston 
Public Library last winter. That so much concerning the habits 
of this insect had been determined at such an early date in the 
history of American economic entomology, is indeed surprising. 
In fact, almost all of the information concerning the insect 
in America dates back to the account of Peck. 

The insect is widely distributed, having been recorded in 
Europe, North America, Australia, New Zealand and South 


Generations. There are in central Iowa two quite distinct 
generations of the insect. In the fall a very few slugs were 
found, lagging way behind the others, but these are probably 
only delayed individuals from the second generation, rather than 
a partial third generation. 

About Ames the eggs and adults were found in late May and 
early June and larvae appeared in the fore part of June. By 
July 5 the first brood slugs were practically all gone. The 
adults from this brood of slugs appear about the middle of July 
and their eggs hatch late in the month. The slugs begin to 
mature a little before the middle of August and by September 
1st they are practically all gone. 

A part of the first brood larvae do not complete their trans- 
formations at once after entering the soil but remain as larvae 
until the next spring, making but one generation a year for a 
part of the slugs. This was noticed by Professor Peck and later 
by Marlatt. It seems to be a provision for the preservation of 
the species. Should the second generation be wiped out by 
natural enemies, the slugs that are held over would still be left 
to propagate the species the next season. 

The Egg. The eggs are deposited in the leaf tissue of the 
various food plants — placed just beneath the upper epidermis 
and thrust through the leaf from below. 

In the insectary eggs hatched in 10 to 14 days in the spring; 
average, 12.1 days; in summer in 7 to 13 days, average, 10.5 

1911] Notes on the Pear-Slug 183 

Parthenogensis probably occurs with this insect, but this has 
not been satisfactorily proved. Many sawflies were examined, 
but no males were found. Eggs deposited by virgin females 
hatched, but all the larvae were weak, and none even reached the 
second stage. 

The Larva. The newly hatched slug is about 1.2 mm. long, 
at first free from slime, but this is secreted in a few hours. The 
young slugs begin to eat out tiny patches in the epidermis; 
later they eat through the parenchyma to the lower epidermis, 
leaving the veins. One slug ate a space of 825.9 square milli- 
meters during its lifetime, 19 days, according to a record kept 
by Mr. T. M. McCall, insectary assistant at the time. 

After each molt, except the last, the larvae eat their cast 
skin, all of it but the head. 

Five molts are probably normal. In 14 cases 5 larvae 
molted 5 times; 5, 6 times; 3, 7 times, and 1 larva molted 8 
times, all under insectary conditions. Sometimes in succes- 
sive molts the width of the cast head was exactly the same; 
often there was very little difference. 

The average head widths of the stages were: Stage I, 
.35 mm.; II, .51 mm.; Ill, .67 mm.; IV, .85 mm.; V, 1.04 mm.; 
VI, 1.13 mm. The geometrical ratio between the successive 
head widths was practically .35. 

Larvae matured in 13 to 26 days; average 19.4 days. No 
essential differences were noticed in the two generations of the 
time required; the same total length was found in either case. 
The time required for the separate stages was: I, 5 days; II, 
3.2 days; III, 3.4 days; IV, 3.2 days; V, 4.7 days (average from 
both generations). Stage VI usually entered the soil imme- 
diately after molting. 

Both Peck and Marlatt remarked that heavy rains are said 
to destroy the slugs. Professor Peck, however, said that he had 
seen the slugs retreat to the under sides of the leaves in a shower ; 
a fact that we observed several times in the Iowa work. It 
seems quite likely that the supposed efficiency of heavy rains 
is really only the disappearance of the slugs to the lower sides 
of the leaves. 

How the larvae reached the ground was a puzzle to us at 
first, but we found that they dropped down of their own accord. 
Occasionally orange-colored larvae were found on tree trunks, but 
they were not crawling down, but appeared to be going upward. 

184 Annals Entomological Society of America [Vol. IV^ 

Finally some black cloth screens were placed on the ground under 
infested cherry trees and the larvae dropped readily to the 
cloth. They were most active about 4 o'clock in the afternoon. 
Eighteen larvae dropped to cloth screens between 2.15 and 4 
p. m., June 29, according to notes made by Mr. T. M. McCall. 

The Cocoon and Pupa Stage. The small earthen cells 
of the larvae were found commonly in the summer within an 
inch of the surface of the soil. Probably they go deeper for 
the winter. 

In the summer it was about 20 days after the slugs entered 
the soil that the adults emerged. In 19 cases the average time 
was 19.9 days, with a range of 15 to 23 days. 

The Adult. The sawflies were most numerous early in the 
morning, although they were found in small numbers at other 
times in the day. In July the sawflies all appeared at about the 
same time, but in the spring the emergence was spread over a 
much longer time. 


Egg Parasites. The most common egg parasite was 
Pentarthron minutum Riley, kindly determined for me by Mr. 
A. A. Girault. It is a very tiny, yellow-brown species, which 
has often been reared from the eggs of a number of common 

At Ames this species was reared abundantly in 1909, but 
not at all in 1910. A few individuals were reared from eggs 
collected at Reinbeck, Iowa, in August, 1910. 

From the first brood eggs the parasites emerged from June 
2-8 to July 20; from the second brood eggs, from August 11 to 
August 20th. 

The eggs affected by this parasite turn black, and of course 
fail to hatch. Two or three individuals were reared from a 
single egg. 

Professor Peck in his original account mentions what is 
doubtless this same species and gives figures of it. After look- 
ing over the account I came to the conclusion that Peck had 
reared the same parasite in Massachusetts in 1798 that I 
reared in Iowa a hundred years later. Mr. A. A. Girault, to 
whom I referred a copy of this account, corroborated my opinion 
that Peck's parasites were most probably Pentarthron minutum. 

1911] Notes OH the Pear-Sing 185 

The second egg parasite, Closterocerns ciuctipennis Ashmead, 
also determined by Girault, was reared equally abundant in 
1909 and 1910, from eggs collected at Ames. This species 
also affected both broods of eggs, emerging from first brood 
eggs June 28 to July 22, and from second brood eggs August -l 
to August 21. 

This parasite was always reared singly from the eggs. The 
black form of the parasitic pupa could often be distinguished 
within the parasitized eggs. It is presumably a primary 

Predaceous Enemies. In 1910 nymphs and adults of 
Podisus maculiventris Say were found to be very active against 
the slugs. Most frequently the slugs themselves were attacked, 
but the Podisus also captured the adults, according to observa- 
tions made by Mr. T. M. McCall. 

One Podisus nymph was kept 47 days in an insectary cage, 
during which time it ate 6G slugs; 1.4 slugs a day. The great- 
est activity was immediately after the insect became an a'dult, 
when it ate 6 and 7 slugs a day. 

A species of Chrysopa and an undetermined Reduviid were 
also observed to prey on the slugs. 

experiments with insecticides. 

Paris green, 1 pound to 150 gallons of water, was effective. 
A home-made arsenate of lead, single strength, was not effect- 
ive. Used against the older slugs this spray was very ineffi- 
cient. I could not see that any slugs at all were killed by it. 
When used against the very young slugs, about half of them 
were killed. 

A 10% solution of kerosene emulsion appeared to be quite 

Several simple soap solutions were found effective, at con- 
centrated strengths. Whale oil soap, 1 pound in 2 gallons of 
water; Ivory soap, 1 bar in 2 gallons and White Laundry soap, 
1 bar in 2 gallons of water, were all satisfactory treatments. 
These soap solutions may be quite convenient for use on a few 
cherry trees when the fruit is present. 

186 Annals Entomological Society of America [Vol. IV, 


A Member: I should like to ask if it is possible to use the 
spraying that is applied for curculio to kill the slug, or whether 
the slug works so late in the season that poison applied for the 
curculio would be washed from the leaves. 

Mr. Webster: The spraying should be done about the 
middle of June in central Iowa. I am not familiar enough with 
the curculio to say whether this spraying would affect it or not. 

A Member: It would be too late to arrest the curculio. 

Dr. Howard : I regret that I did not hear all of the paper. 
I should like to ask about the affect of a strong stream of water 
on the slugs, without any insecticide at all. 

Mr. Webster: I did not try that. 

Dr. Howard : In city yards a strong stream of water from 
a hose can be used to wash off the slugs. This method has been 
tried and given good results. 


By Henry P. Severin, Ph. D., Professor of Zoology and Entomology, 

College of Hawaii, and Harry C. Severin, M. A., Professor of 

Entomology, South Dakota State College of Agriculture 

and Mechanic Arts. 

(with plate XIV.) 

In the Phasmidae, Mantidae, Blattidae and Acridiidae, the 
cervical ampulla is said to play an inportant role in the process 
of molting, and in some Orthoptera, also in the process of hatch- 
ing. This ampulla, consisting of a soft membrane joining the 
head dorsally to the prothorax, can be transformed by the afflux 
of blood into a greatly swollen pouch, which then projects out 
immediately behind the head. 

The process of hatching of various Orthoptera has been 
studied by a number of entomologists. Riley (7) does not men- 
tion the cervical ampulla while describing the phenomenon of 
hatching in the Rocky Mountain Locust, for he writes as fol- 
lows : "The hatching consists of a continued series of undulating 
contractions and expansions of the several joints of the body, 
and with this motion there is slight but constant friction of the 
tips of the jaws and of the sharp tips of the hind tibial spines, 
as also of the tarsal claws of all the legs against the shell, which 
eventually weakens and finally gives away. It then easily 
splits up to the eyes or beyond, by the swelling of the head." 

Packard (5) objects to Riley's account of the supposed 
action of the jaws and spines and believes that "the egg-shell 
is without doubt burst open by the puffing out or expansion 
of the membrane connecting the head and prothorax, just as the 
common house-fly or flesh-fly bursts off the end of its pupa- 
case by the puffing out of the front of the head." 

Kunckel d'Herculais (3 and 4) gives the following account 
of the physiological mechanism in the hatching of the Acridii- 
dae: " Les Acridiens rompent la coque de I'oeuf, * * * par la pres- 
sion exercee a I'aide de la membrane unissant dorsalement la 
tete au prothorax que se transforme par afflux de sang en une 
ampoule cervicale." 

In Diapheromera femorata the mechanism, which ruptures 
the various membranes and springs off the operculum when the 
walking-stick is about to emerge from the egg, cannot be ob- 
served in action on account of the hard, thick, opaque chorion. 
If the operculum is carefully removed from an egg shortly 


188 Annals Entomological Society of America [Vol. IV, 

before hatching, the embryo will be found with its head and pro- 
thorax situated directly beneath the portion of the egg removed 
(Fig. 1, h and p). The pressure exerted by the cervical ampulla 
is, therefore, directly against the operculum. 

Hatching spines for the purpose of rupturing the embryonic 
envelopes and also for breaking or cutting open the egg-shell 
have been described from many insect eggs. Above the pro- 
thorax of Diapheromera, the thin amnion is covered by num- 
erous long spines which point toward the operculum. These 
spines, like the egg-burster (or ruptor ovi as Riley (6) calls it) 
of Corydalus cornutus, are portions of the amnion itself. If 
the prothorax of a walking-stick is examined after its emergence 
from the egg, no spines are found, but simply short blunt pro- 
tuberances. In all probability, the long spines of the amnion 
above the prothorax assist in rupturing the vitelline membrane 
which is especially thick beneath the operculum. 

"When the young walking-stick is in the egg, ready to emerge, 
the meso- and metathorax are not remarkably elongate, but 
before the little creature is fairly out of its narrow prison, the 
thoracic segments assume their usual proportions. It is said 
to be a most curious sight by those who have observed this 
almost instantaneous development." (Caudell [2]). 

An attempt was made by us to secure an explanation for 
this curious phenomenon observed by Caudell. After the 
chorion of the egg was removed, the embryo was found to be so 
curled up in the egg that the posterior end of the abdomen lay 
near the head region. A longitudinal section through the 
embryo showed that the thorax was folded transversely in a 
dorso-ventral direction (Fig. 2). In all probability it is simply 
the straightening out of these folds as the young walking- 
stick emerges, that causes the thoracic segments to assume their 
usual proportions. If the pressure exerted by the cervical 
ampulla at the time of hatching is not sufficient to rupture the 
amniotic and vitelline membranes and also to throw oflf the 
operculum, it may be possible that the straightening out of some 
of these thoracic folds assist in the process. 

After pushing off the operculum, the young walking-stick, 
with the prothorax bent down at its union with the mesothorax, 
begins to emerge from the egg (Fig. 3). The cervical ampulla 
is now slightly swollen, and the prothorax possesses a deep 
green color, due to the blood which has accumulated within it. 


Hatching of the Walking Stick. 


The method employed during the process of emerging from 
the egg is almost identical with that which we (10) have des- 
cribed of a walking-stick withdrawing itself from its old skin 
during the process of ecdysis. A specimen examined under a 
binocular microscope during the process of emergence from the 
egg, will be seen to undergo a series of peristaltic-like movements 
of the segments of the body; these movements pass from the 
posterior end of the abdomen towards the head. With each 
series of these movements, the body is drawn out of the egg- 
shell a short distance, the legs also assisting somewhat in this 
process of extraction. At each pull of the legs in their attempted 
withdrawal from the egg-capsule, the strength of the pull is 
such, that the coxa of each leg presses against the body, causing 
in that region a temporary indentation. When the peristaltic- 
like movements reach the head, the walking-stick often raises 
the head vigorously upward in an attempt to withdraw the 

The first part of the walking-stick to leave the egg is the 
dorsal surface of the prothorax (Fig. 3, p) ; then comes the head 
(Fig. 3, h), followed by the rest of the thorax. The antennae 
are freed next, and these may come forth either simultaneously 
or one soon followed by the other. The following order was 
often observed in the withdrawal of the legs: one middle leg 
was followed by the other; then the front legs were pulled out 
of the egg at the same time,' and finally the hind legs. The 
abdomen does not leave the egg at any definite time in relation 
to the withdrawal of the other parts, but it may emerge after the 
antennae or, in other specimens, after the middle or front legs. 
The extrication of the antennae, legs and abdomen, however, 
does not always take place in the order just given, as is shown 
in the following table: 

Table I. 

Order of Withdrawal of the Antennae. Legs and Abdomen from the Eggs of Six 
Diapheromera femorata. 

front leg 
middle leg 
middle leg 
front leg 
/hind leg 
\hind leg 

middle leg 
middle leg 
front leg 
front leg 
fhind leg 
\hind leg 

middle leg 
middle leg 
front leg 
front leg 
hind leg 
hind leg 

middle leg 
middle leg 
front leg 
front leg 
fhind leg 
\bind leg 


middle leg 

front leg 

front leg 
(middle leg 
\hind leg 

hind leg 

thrown ofT. 

middle leg 
middle leg 
front leg 
front leg 
hind leg 
hind leg 

Braces indicate that the two included appendages were extricated simultaneously. 

190 Annals Entomological Society of America [Vol. IV, 

Stockard (11) describes the hatching of Aplopus mayeri as 
follows: "When hatching the embryo's head and body come 
forth from the egg first, the antennae are then pulled out, the 
legs being the last parts liberated from the shell." The speci- 
mens noted under E and F in the above table agree with Stock- 
ard's observations on Aplopus, but both of these specimens 
had their appendages caught in the amniotic membrane (Fig. 
4). In a previous paper we (9) have already called attention 
to the fact that dryness, at the time of hatching, has a marked 
effect upon the emergence of the walking-stick from the egg. 
With the addition of water which was added drop by drop to the 
egg-shell, within which the above-mentioned specimens were 
caught, these walking-sticks succeeded in freeing themselves. 


1. Bourgeois. J., 1900. Sur un mode particulier de progression souterraine chez 

quelqucs larves d'Insectes. Bull. Soc. ent. Fr. LXVIV, pp. 261-2. 

2. Caudell, A. N., 1903. The Phasmidae, or Walking-Sticks of the United 

States. Proc. U. S. Nat. Mus. XXVI, No. 13.3."), p. 864. 

3. Kunckel d'Herculais, J., 1890. Mecanisme physiologiques de Teclosion, 

des mues et de la metamorphose chez les Insectes Orthopteres de la famille 
des Acridides. C. R. Acad. Sci. Paris, CX, pp. 657-9. 

4. , 1890. Du role de I'air dans le mecanisme physiologique de I'eclo- 

sion, des mues et de la metamorphose chez les Insectes Orthopteres de la 
famille Acridides. Ibid. CX, pp. 807-9. 

.5. Packard, A. S., 1877. 9th Ann. Rept. U. S. Geol. and Geographical Survey, 
pp, 630-4. 

6. , 1903. A Textbook of Entomology, pp. 585-6. 

7. Riley, C. V., 1877. 9th Ann. Rept. Noxious, Beneficial and other Insects of 

the State of Mo., pp. 89-90. 

8. Riley, C. V., Packard, A. S., and Thomas, C, 1878. 1st .\nn. Rept. U. S. Ent. 

Com. 1877. Relating to the Rockv Mountain Locust. Wash. pp. 277-284. 

9. Severin, H. H. P., and Severin, H. C, 1910. The Effect of Moisture and Dry- 

ness on the Emergence from the Egg of the Walking-Stick, Diapheromera 
femorata Say. Jour. Econ. Ent. Ill, No. 6, pp. 479-481. 

10. , 1911. The Life-History of the Walking-Stick, Diapheromera fem- 
orata Say. Jour. Econ. Ent. IV, No. 3. 

11. Stockard, C. R.', 1908. Habits, Reactions and Mating Instincts of the Walk- 

ing-Stick, Aplopus mayeri. Publ. No. 103, Carnegie Inst., Wash., pp. 

All figures were drawn with a camera lucida. 
Fig. 1. View of embryo after the operculum has been removed showing the 
head and prothorax directly beneath. The pressure exerted by the cervical 
ampulla, which joins the head dorsally to the prothorax, would be directly 
against the operculum: h, head; p, prothorax; vi, vitelline membrane; c, compound 

Fig. 2. Longitudinal section through the head and thorax of the embryo, 
showing the transverse folding of the thorax in a dorso-ventral direction; h, head; 
p, prothorax; ni, mesothorax; met, metathorax. 

Fig. 3. Walking-stick emerging from the egg, showing that the prothorax 
is bent down at its union with the mesothorax: h, head; p, prothorax; m, meso- 
thorax; op, operculum still adhering to the egg by means of the so-called "shell 

Fig. 4. Walking-stick with its appendages caught within the egg-shell: 
ha, "hilar area." 

Annals E. S. A. 

Vol. IV, Plate XIV. 

H. H. P. and H. C. Sen 



Many j^ears ago in England, I captured a rather uncommon 
hemipterous insect, and sent a record of it to a well-known 
entomological journal. The editor, being a lepidopterist, had 
never heard of the bug, but did know of a very rare moth hav- 
ing the specific name {bicolor) employed. He accordingly 
changed the generic name to that of the moth, and I found 
myself the astonished recorder of an insect I had never seen 
alive, nor hoped to see. More recently I communicated to a 
publication in this country a short paper on a supposed new 
plant of the genus Ribes. The editor, not liking the title, 
substituted "A New Currant from Arizona," whereas the plant 
was a gooseberry, and was from New Mexico. These rather 
amusing instances are cited merely to illustrate the indisputable 
fact that it is risky for an editor to interfere with the contri- 
butions he publishes. On the other hand,. I have been shown 
manuscripts sent in for publication which, if printed exactly 
as received, would be simply unintelligible. The editor is in a 
difficult position, and as a rule, I think the contributors have 
little reason to feel otherwise than grateful for the treatment 
they receive; it is at least not rarely better than they deserve. 

Although I am against editorial alterations in manuscripts, 
I think it may be entirely proper to adopt some simple rules to 
be enforced in every case, the papers which fail to conform 
being returned to their authors for correction. As entomologi- 
cal editors appear to have no such rules, with the exception of 
a few relating to typography, it occurs to me that the Entomo- 
logical Society might properly discuss and adopt a set, pressing 
them upon the attention of editors with such authority as it 
may be considered to possess. As the result of a little private 
correspondence, I believe it would be easier to get all the editors 
together to agree upon certain things, than to persuade them 
individually to take the desired step. I cannot do more than 
present a suggestive^^ outline, which may be discussed and 
amended as necessary. 

(1) When a new genus is described, the type species must be 
stated; it may be as well to add, that the binomial made by 
combining the generic name with the specific name of the type 
species must be printed. 


[1911] Entomological Publications. 193 

(2) No new genus will be published, that is not based on a 
described species. 

(3) Rules 1 and 2 also apply to subgenera. 

(4) No new species may be described without comparing it 
with some other described species, or stating wherein it differs 
from other members of the genus. 

(5) When a new species is based on specimens from several 
localities, it must be explicitly stated which is the type locality. 

(6) When a new species is described the data concerning 
localities and collectors must be given in full so far as known. 
This is also strongly recommended in the case of all new records. 
If the locality, collector, etc., are not known, it may be well to 
say so, although this may be taken for granted if the writer is 
known to be careful in citing data. 

(7) It is impossible to avoid all errors in spelling, grammar, 
etc., but so many of them have appeared in recent years, that 
American entomologists have some reason to feel ashamed. 
It would be easy to compile a list of scientific names which must 
be retained in our lists, although faulty to the extent of being 
offensive. This is true in spite of the freest recognition of the 
fact that scientific latin is a living and growing language, and 
must include many words unknown to the ancients. No rule 
can cover this difficulty, but it might be worth while to collect 
every year a list of these criticisable productions, and set them 
forth as a warning to authors and editors alike. 

(8) It is not permitted to publish new varieties as binomials; 
the trinomial must in every case be written out. 


By Richard A. Muttkowski. 


I. Standards for Descriptions. 

A. Specific description. 

B. Redescription. 

C. Generic description. 

II. Standards for Colors. 

III. Standards of Nomenclature. 

A. Generic and specific nomenclature. 

B. Anatomical (morphological) nomenclature. 

C. Nomenclature of types. 

IV. Standards for Keys (tables) of Genera and Species. 

V. Standards for Indices. 

VI. Standards for Titles. 

VII. Standards for Reprints. 

Taxonomy is the most important incident of Science — 
things must be named before we can write of them. If we 
consider the proportion of zoological papers at the present time, 
we find that about two thirds are systematic. But this pro- 
portion increases to seven-eighths if entomological papers 
alone are considered. 

In taxonom}^ we usually speak of only two types of contri- 
butions: (a) text books, prepared for a mixed public, and ih) 
original work, which is intended for the entire scientific world, 
but whose chief appeal is directed to a small group of contribu- 
tors. These form a special class, distinct in conception and 
treatment from other scientific papers. 

The style of composition proper in a morphological, onto- 
genetical or phylogenetical paper would be found impracticable 
in a taxonomic paper. Yet, while the greater number of 
entomological papers are taxonomic, it nevertheless remains a 
curious fact that as yet no compendium for taxonomic comj^o- 
sition has been published. 

For papers other than taxonomic we have Dr. T. Clifford 
Albutt's excellent book, "Notes on the Composition of Scien- 
tific Papers" (MacMillan Co., New York, 1904, 8vo). While 
many of the chapters of this work would be of interest to syste- 
matists, they do not pertain directly to taxonomy; the volume, 
on the whole covers a quite different ground, that of scientific 


1911] Composition of Taxonomic Papers 195 

To say that nothing at all has been published on the present 
issues would invite criticism. On the contrary, I have found 
copious and more than sufficient material in the more prominent 
journals of the past two decades. But these contributions are 
widely scattered; moreover, they are written as protests and 
usually deal with a single topic only. While I cannot lay claim 
to originality in the suggestions included in this paper, I have 
endeavored to treat all of the more vital topics bound up with 
taxonomy ; aiming to suggest such standards in writing as would 
conform to the various needs of those interested. 

Briefly stated, a standard is the result of an average or con- 
sensus of opinions upon a given subject, hence a criterion. 
Thus far the only criterion of any worker has been the approval 
of his fellow- workers along the special line of work he has adopt- 
ed. This basis is hardly sufficient, as not a small coterie of 
workers but the world at large is intended to be benefited. 

It must be remembered that this paper does not treat of 
criterions of species, but with the composition of descriptions 
and general methods of presentation. That these are perfect 
and above reproach probably none will maintain. 

I have talked over these matters repeatedly with scientific 
workers. Curious to say, dissatisfaction with present methods 
and with the absence of definite standards was prevalent every- 
where. The necessity of co-operation toward the achievement 
of practical standards was sometimes very strongly expressed. 
A digest of all these opinions, private or published, may be 
summed up in the following: Better methods of description are 
desirable; the nomenclature of species and genera, of colors and 
types to be regulated; titles of articles to be made more com- 
prehensive; reprints to contain place, time and name of publi- 
cation, etc. 

I. Standards for Descriptions. 

A. Specific Description. — J s , an unencumbered species: Front 

pale, palpi scaled, thorax with black stripes, antennae yellowish, abdomen spot- 
ted, legs with pale, wings with black markings. Head with short pile, abdomen 
tufted, a small tuft between the antennae, which are fuscous at the base, white 
toward the tips. Wings white, with four or seven black lines crossing them, 
the lines curved or straight. Abdomen with tufts black, exceeding anal angle. 
Legs long, with spurs. Palpi reaching the vertex, legs slightly darker at the joints. 

I dare say, that no living man could determine a specimen 
from this extravaganza. Yet the description is made from an 
actual sYieciQ^ — Conchylodes platinalis, Lepidoptera-Pyralidae. 

196 Annals Entomological Society of America [Vol. IV, 

Furthermore, it is typical of many descriptions of the past, and, 
I regret to say, of too many in the present days. 

It is curious how little logic is often applied in formulating 
a description. One specialist, who has written hundreds of 
descriptions, seems to have found particular pride in making 
these as intricate and involved as possible; there is no logical 
sequence in the treatment of the main divisions; on the contrary', 
the acrobatic description jumps from antennae to legs, from 
abdomen to head, wings to palpi, venation to tarsi, morpholo- 
gical characters to vestiture, color to structure, etc., etc. So 
much so, that after comparison is completed one must begin 
over again, as it is impossible to remember the way through 
the labj'rinth. 

Descriptions should not be written for personal aggrandize- 
ment, but to announce a new fact or discovery to the scientific 
world. Such being the case, the description, once published, 
belongs to the world at large and no longer to the writer. The 
author therefore owes it to science that the facts of which the 
world is to become owner be presented in a manner most acces- 
sible to, and best applicable by other men. If the author for 
any reason whatsoever is careless and inaccurate he sins against 
science. After all, there is an intellectual as well as a moral 

"Head pale, eyes small, dark, vestitute smooth and yellowish, 
body moderate, legs short, tibiae stout," applies equally well 
to Mr. Jones as to Pediculus capitis strolling on his head. 
Brevity may be the point of wit, but science is no joke; taxonomy 
deals with facts, not idiosj^ncracies. Who has not felt the bane 
of two to eight lined descriptions, any one of which harmonizes 
easily with half a dozen or more distinct species? I do not be- 
lieve that an entomologist lives who has not at one time or other 
execrated these brief, vacillating descriptions. But why do ento- 
mologists continually write others that are no whit better or 
longer? Let it be known, that one thorough description cover- 
ing three pages may be of more use and more valuable to sci- 
ence than three descriptions on one page. One may suggest that 
the perusal of brief descriptions saves time; but when we come 
to analytic comparison of closely related species the brief descrip- 
tion forms an obstacle which results in considerable loss of time. 
What of genera whose species are extremely variable? Can the 
extent of specific variation together with a description of the aver- 

1911] Composition of Taxonomic Papers 197 

age be summarized in twenty lines? Hardly. The ideal descrip- 
tion will be a careful analysis of all body parts with all their 
appendages, attributes and characteristics, to be followed by a 
summary of salient characters of the type and a comparison to 
related species. I maintain that this cannot be accomplished 
on less than a page. 

It is terrifying and discouraging to be confronted by a page 
of solid description, where all characters, whether head, thorax, 
abdomen, or wings, flow together in a solid phalanx, so that it is 
impossible to pick out readily any special point desired. Des- 
criptions should be paragraphed or captioned. This costs no 
extra labor, and, in fact, presents a much neater appearance 
when published than the solid, uniform mass of words. Besides 
it affords greater facility to the student who wishes to look up 
certain characters for comparison. 

Again, a description should not be isolated. I mean, com- 
parison to related species and indication of the position of the 
new species should follow the description. It is reprehensible 
negligence to describe a new species from a genus already con- 
taining a dozen or more species and to omit all mention of either 
relations or position ; such proceeding is indeed worthy of repri- 
mand. To say the least, the work of the author will be placed 
in an extremely doubtful light. The thought suggests itself, 
that the author himself was ignorant of the relations and that 
he described a species at hap-hazard. 

After all this, why pay any attention to identity, number, 
and custody of types? Why state the locality from which the 
types came? Why select a holotype from a series of twenty 
specimens that show considerable variation? No one is ever 
expected to express any doubt of the scientific determination of 
the twenty. No one is ever expected to feel interested in looking 
up the types for comparison or study after having become famil- 
iar with the all-sufficient description of ten lines. This seems 
to be the opinion of some taxonomists. For they very carefully 
avoid all mention of the number of types, their identity (see 
nomenclature of types) and only grudgingly designate the locality 
from which the types came by the remarkably precise state name. 
The latter, it is supposed, will give the reader all the ethological 
information he desires ; so that if he wishes to capture specimens 
of the same species, all he need do is to pack his trunks and hie 
himself to "Texas" or "Nevada" and pick the species from the 

198 Annals Entomological Society of America [Vol. IV, 

mountain-sides and valleys, from water and land, from trees 
and grass, or just open his bottles to stop their fall from the 
heavens. It must be there, for the author said so; he said 
"Nevada" and this is Nevada. 

The following is a scheme for an accessible description: 

J s . Not a new species: 

1. (a) Sex, usually cf , and dominant color; size. 

(b) Head: mouthparts, face, eyes, vertex, antennae, occiput, etc.; vestiture, 

colors, stnicture, etc. 

(c) Thorax: prothorax, mesothorax, metathorax, structure, vestiture, colors. 

Legs, their color, structure, vestiture and appendages; etc. 

(d) Abdomen: structure, markings, color, vestiture, appendages, etc. 

(e) Wings: color, markings, vestitute, venation, etc. 

2. (a) 9 and dominant color; size. 

(b, c, d, e) as above. Difference from d'. 

3. Summary of salient characters. Unique characters. Variation. 

4. Comparison to related species, position of species. 

5. Material: Tj'pes, identity (see nomenclature) of types, exact date and locality 

of capture. How (ethology) and by whom captured. 

Of course, this scheme cannot be strictly adhered to in the 
different orders; it is, however, sufficiently elastic to permit the 
changes required. What is important in one order, is negligible 
in another. But the fundamental idea of setting forth by para- 
graphs or captions the principal parts of a description in suc- 
cessive order, will no doubt be understood. 
- B. Redescription. — How a redescription should be formed 
depends on the original description. If the original was care- 
fully drawn, the other may be a summary of the first with possible 
new points of variation, etc., discovered. Or if, as very often 
is the case, the original was insufficient, the redescription should 
be carefully formed; in fact, the author should aim to replace 
the first with the second description. Even though his name 
stand not as the sponsor of the species, the task of redescribing 
is not a thankless one, as need hardly be explained. 

Redescriptions are also written for convenience, either as 
summarizing the knowledge of the species, or, as indeed com- 
mendable, to place an otherwise inaccessible description within 
the reach of the student. Much of what has been said under 
the preceding caption applies here also and needs no repetition. 

C. Generic description. — What is a genus? A classifica- 
tory group of plants or animals, embracing one or more species; 
the primary condition of binary nomenclature; a uninominal 
used for the lowest phase of the grouping of living forms ac- 
cepted by naturalists. 

1911] Composition of Taxonomic Papers 199 

What constitutes a genus? A single species or several that, 
aside of specific dififerences, have certain morphological features 
in common, which distinguish them from all other groups of 

When is a genus valid? When so stated by the sponsor, 
the nomenclator having noted certain morphological characters, 
the value of which is recognized by fellow-workers, and who 
accept this diagnosis upon the given characters; when placed 
with a monomial (specific) to signify that the species possesses 
certain distinguishing group characters. 

As genera constitute the lowest, but at the same time the 
most important, phase of grouping, at least some attention 
should be given to the formation of generic descriptions ; espec- 
ially so in larger contributions, such as monographs and generic 
summaries and synopses. Generic description is allied to speci- 
fic description; hence methods ought to be similar. 

Some of the essentials of a generic descrijjtion are the 
following : 

1. That the type species be cited. It should be noted that the 
type species must be a species then or previously described; else we 
have merely a nude name. 

2. That the characters on which the species is leased be given. 
Although the generic name alone, when coupled with a described spe- 
cies, is recognized as valid by the codes, the systcmatist will insist that 
the absence of a generic description is an unfair apprisal on the part of 
the nomenclator. 

3. That these characters be stated concisely; that is, write to the 
point. Brevity is not conciseness. One may be brief and vague at 
the same time. 

4. That these characters be stated in orderly manner. Especially 
in larger jjapers unity of methods is advantageous. If one description 
begins with the legs, another with head, a third with the venation, etc., 
study is made difficult. Unifonnity of methods facilitates study and 

5. That other species belonging to the new genus be listed. While 
this necessitates thorough stud}' on the part of the nomenclator, it 
really is his duty. To split up large genera upon characters drawn 
from a single species is a simple matter. But the nomenclator should 
verify the stability of his characters by extensive comparison with 
related species. 

6. That other genera be compared, or, at least, the position of the 
new genus indicated. To describe a new genus of a family already 
containing twenty or thirty genera and not indicate the position or 
relations of the newcomer, is not scientific; it denotes carelessness or 

200 Annals Entomological Society of America [Vol. IV, 

Not exactly essential, but still of value in extended papers 
are the following: 

7. The etymology of a new name should be explained. This 
often gives a clue to a character or to the relation of a genus. 

8. The distribution of a genus should be cited; namely, whether 
it is Oriental, Palearctic, Nearctic, etc. 

9. The order and family of the genus should be indicated in title 
or text. This pertains especially to brief papers. As nobody can be 
familiar with the specialties of all authors or with all generic and family 
names, this offers an aid in the study and classification of papers. 

10. The validity of the generic name should be considered by the 
author. It is the author's province to do so in the first place. If 
taxonomists cared to put a little time or expense to the verification of a 
name, there would be fewer homonymns coined each year. 

II. Standards for Colors. 

Of all standards these are most needed, since they are most 
sinned against. That no color standard should exist in a divi- 
sion of zoology, which is of prime importance economically as 
well as numerically, and where frequently colors are our only 
tangible gixides — unfortunately so — for -generic and specific 
determination, is hardly conceivable. Yet such is the case. 
After one and a half centuries of entomology, in which the 
number of described species has been advanced from a few 
hundred to several hundred thousands, we are utterly lacking of 
any color standard and are guided in our nomenclature of col- 
ors solely by the individual impressions of the taxonomist. 
That sttch a basis is absolutely at faidt, needs no special 

The perceptions of most men in regard to colors are extremel}^ 
crude. (To anyone who may doubt this statement I advise a 
visit to some artist. One may state to him his impressions of 
ten different shades of color; and observe then, how often the 
shade will be misnamed by the amateur as against the profes- 
sional testimony of the artist. I do not claim a better perception 
than other men and am found at fault equall}^ as much as others.) 
In their school days men were taught the tale of three to seven 
primary colors, and a small trifle of the shades resulting from 
combinations of the primaries. A little of this they remember 
throixgh the rest of their lives. And, strange to say, when a 
man would not use a term or expression to designate an anatomi- 
cal detail unless he is absoltitely certain that it is correct, this 
same man will unhesitatingly designate colors, when, to say the 

1911] Composition of Taxoiiomic Papers 201 

least, there is good reason to doubt his exact knowledge of the 
particular color. I do not say that this is intentional ; it results 
from overconfidence of his particular knowledge. This care- 
lessness arises from the lack of proper standards. Accordingly 
men are forced to formulate their own standards, which are 
necessarily at fault It is only through an average or consensus 
of opinions that standards are reached. 

In a desire to be conscientious men often circumscribe a con- 
dition when they find their exact knowledge of colors inadequate. 
This is usually done by the addition of such terms as "pale, 
light, medium, shining, glabrous, bright, vivid, dark, dull," 
etc., to the primary color. While this effort is commendable, 
it offers no more certainly than the mere citation of the primary 
shade; and the interpretation of the circumscriptiye adjective 
is frequently very liberal. 

Probably the most liberty has been taken with the term 
"fuscous" in our descriptions. This term has been made to 
designate any darker shading on a light back-ground, begin- 
ning with a tinge of the palest yellow against a white or trans- 
lucent base to a seal or clove brown against any lighter back- 
ground. "Orange," "yellow," and "green" are others of these 
liberally interpreted colors. The heart-rending or laughable 
(as one views it) puzzling of students, who are familiar with 
exact anatomy but not with the vagaries of taxonomy, when 
attempting to determine a species from description and to seek 
conformity between the colors as given by the author and the 
specimen in hand, affords too well known illustration. 

Viewing the matter from the stand-point of my own desul- 
tory experiences, the question occurs to me: If at the present 
time, when the approximate number of described insects 
amounts to about 300,000 species, identification is difficult, 
the determination often exhausting the patience of the taxo- 
nomist in the vain endeavor to divine the protologist's percep- 
tions of colors; further, this difficulty having encumbered tax- 
onomy with labyrinthine synonymy; — what, then, will be the 
condition of taxonomy fifty years hence, if we continue with 
present methods, when species will have increased to approxi- 
mately 1,000,000? 

Happily there is a tendency among our eminent specialists in 
the last decade to standardize their descriptions as far as colors, 
are concerned. (This is beautifully instanced by Packard in 

202 Annals Entomological Society of America [Vol. IV, 

Tiis later works, such as his monograph of the Lepidopterous 
family Notodontidae). Yet these are so few that their number 
may be regarded as negligible. That the necessity of color 
standardization is imperative and that this is well recognized 
is shown by Dr. J. B. Smith's addition of a plate of colors to his 
recent "Dictionary of Entomological Terms." 

Structural (iridescent) colors are sometimes difificult to 
define because of the varying hue, according to the angle of 
refraction and reflection. Yet with reliable color charts these 
difficulties would be obviated. 

Frequently the belief asserts itself that specimens were 
described in lamp-light. How unsatisfactory and misleading 
artificial light is taxonomists ought to know only too well. 
The simple experiment of exposing green, yellow and brown 
insects, notably shining specimens, successively to gas, electric, 
acetylene, candle, kerosene and the natural sun-light yields 
some surprising results. 

A color standard need not be an assortment of infinitesimal 
shadings, gradings, and combinations of the primaries. A repre- 
sentative selection of from thirty to fifty colors is sufficient for 
all practical purposes. 

The fact that detailed comparison of the colors of a specimen 
to color charts entails some extra labor should not deter taxo- 
nomists from making these comparisons. The appreciation 
and gratitude of their fellow-workers as well as of their follow- 
ers will be their reward. The dominant color should be stated 
in all cases. True, the colors of dead insects are rarelj^ quite 
the same as in life, or those of younger insects the same as 
those of mature specimens. Yet the fact that colors have faded 
in death, or that they change with age, is of secondary import- 
ance. A description is not based on possibilities, but on tangible 
concrete actualities. These alone should rule. If there are good 
reasons for assuming that the colors of the specimen are not 
representative, this can, and, in fact, should be stated. Hav- 
ing a dominant color as a basis, it is comparatively simple 
to fix the position, extent, and shade of the other colors an 
insect may exhibit from further comparison to charts. 

The terminology of colors may be somewhat cumbersome. 
But science is not "belle lettres"; the taxonomist does not con- 
sider whether the sentences he reads are syntactically correct 
or rhetorically rounded, but judges from their contents as to 

1911] Composition of Taxonomic Papers 203'' 

their value. At that, why a composite terminology? Why not 
a restricted nomenclature based on a few names with divisions 
indicated by subnumerals, as red 1, red 2, red 3, etc.. blue 1,. 
blue 2, etc., etc.? 

Good works on colors exist, notably Ridgeway's Nomencla- 
ture of Colors, as adopt-ed by Ornithologists. (Unfortunately 
this excellent work is long, out of print, and because of its lim- 
ited edition it is now practically impossible to purchase a copy 
in the book-market). But for practical purposes a simple chart, 
as that hand-painted by Frederick Oughton (London), if se- 
lected by a representative commission of entomologists, could 
be manufactured at low expense, which would be easily justified 
by the demand. This would offer a standard for all times, not 
to mention the other obvious advantages resulting thereby- 

III. Standards of Nomenclature. 

A. Generic and Specific Nomenclature. — This is the only 
sphere where standards already exist. These standards are the 
codes of zoological nomenclature, such as the International 
Code of Zoological Nomenclature, the A. O. U. Code of Nomen- 
clature, etc., which are commonly followed by zoologists. If 
I say "followed by zoologists," the phrase must be given the 
most general and generous interpretation. Speaking of a class 
I can say "commonly" ; but when speaking of groups of special- 
ists — to say it mildly, many groups use the nomenclature of 
1810 instead of 1910. This sounds anomalous, but it is not. 
For the regulation of nomenclature by codes of universal 
sanction is comparatively recent, and the commissions are only 
gradually . bringing order into the nomenclatural chaos that 
existed before their day. 

One cannot expect, I suppose, that a speciahst on the bio- 
logical phase of insects should be interested in the "arbitrary, 
dry" codes of nomenclature. Yet it must be remembered, that 
taxonomists alone have caused the chaos. Taxonomy is' 
"arbitrary" also. What one man considers a variety, another 
calls a distinct species; and still another refuses to recognize- 
either opinion. Or are "spHtters" and "lumpers" only births 
of fancy, or memories of the distant past? 

The aims of the codes of nomenclature are to make the 
nomenclature as free and unencumbered as possible. Hence the 
rules set down for guidance. If taxonomists disdain, or even 
refuse, to follow these rules, who else should follow them? 

204 Annals Entomological Society of America [Vol. IV, 

B. Anatomical {Morphological) Nomenclature. — Standards 
for generic and specific nomenclature have been noted. The 
present issue is of equal significance. 

The chief objection that may be stated on this question is 
indefiniteness. A lesser offense is the scope of the terms; 
e. g., while by "front" the author may intend to include nasus, 
epistoma, rhinarium, labium, etc., we, however, know that 
front means frons in the scientific interpretation and nothing 
else. What the author thinks, we cannot telepathicallj^ or by 
any other means divine. 

Each business has its technical nomenclature. No hardware 
man will hand you a shingling-hammer when you ask for a 
claw-hammer. To the business man the two terms signify 
two different things and he will never be so careless as to use 
the one for the other. Yet among taxonomists we find a con- 
tinual interchange of terms, such as joint for segment, tarsi for 
tarsal claws', mouth for labrum or mandibles, abdomen for ven- 
ter, etc. 

When a taxonomist writes "face yellow, abdomen spotted," 
it is supposed, that he knows what he means. But unfortu- 
nately I do not. A specialist, who knows the peculiarities of 
the score or twenty-five other men working on the same branch 
of science, will possibly understand what is meant. Not so the 
individual who attempts to determine a species, less because of 
special interest, but because of some observation he made on it 
and which he desires to record in his book of field-notes. 

Another idiosyncrac}^ is to use comparative terms for the 
length or size of any portion of the body, as, for example, 
"front as wide as the eyes, elytra twice the width of the pro- 
notum, tarsi about two thirds the length of the tibiae, etc." 
This mode of measurement is miserably uncertain; miserably, 
because of the misery of the student who attempts to make 
the same comparisons and cannot see them as the author 
saw them. 

How many men are able to mark the exact middle of a line 
at a glance? Aside of usual differences in refraction in two eyes, 
some aberration will be caused by the strain of focusing to the 
same point. A "mathematical" eye is a virtue that very few- 
people possess. Still more difficult is to find the exact third of 
a line. What then of paralleling lines, or approaching lines? 
What of curved lines, irregular lines, etc.? Or is the chapter 
on "Optical Illusions" as taught in Physics only an illusion? 

1911] Composition of Taxonomic Papers 205 

Bad as color illusions are, mathematical illusions are worse. 
The chapter on "Optics" ought to form the favorite reading of 
many taxonomists. A difference of one millimeter on an insect 
of 20 mm. length is slight; but it makes a considerable difference 
on an insect of 8 mm. It is a peculair experience to read in a 
description of a beetle or any other insect "elytra twice the 
width of the pronotum" and then find by actual measurement 
that the pronotum is 4 mm. at its widest point while the elytra 
are 10 mm. or more in length. Similarly with most other com- 
parative measurements. When tested by the micrometer or 
millimeter scale they will be found considerably aberrant. 
Hence the urgent advisability to introduce exact measurements 
instead of the unreliable optical method of comparison. 

One standard does exist in anatomical nomenclature, name- 
ly the Comstock-Needham nomenclature of wing venation. 
The merits of this system are undisputed and recognized by all 
modern systematists. But instead of unreservedly adopting 
a system the value of which they confirm, taxonomists inter- 
mingle the antiquated miscellaneous wing nomenclature with 
the logical modern terminology. As a result we are continually 
thrown from one style of naming the veins to the other. This 
may not be troublesome for the specialist. But it a student is 
generally interested in entomology, he finds himself in a constant 
quandary as to the special terminology of each particular order, 
as they are easily confused; whereas the Comstock-Needham 
nomenclature was especially designed to obviate this difficulty. 
It is true, certain orders have certain appendages which it is 
desirable to retain, e. g., for Neuroptera the thyridium cell and 
end-forks, bees the subcostal cells, etc. These should be retained, 
as they are special attributes of the respective order, family or 
genus. But the fundamental principles of venation, as out- 
lined by the Comstock-Needham nomenclature, are possessed 
by all orders, viz., costa, subcosta, radius, media, cubitus and 
anal vein. Why not use them instead of vein 1, 2, 3, 6, 8, 10, 
etc.? The terminology is simpler, it is less aggravating, it is 
more logical, and it is an aid to the student and worker. 

C. Nomenclature of Types. — Quite as important as specific 
and anatomical nomenclature is the nomenclature of types. 
Considerable attention has been given to the latter study in 
recent years. As the various departments of natural history 
are dependent mainly upon descriptions for the taxonomic 

206 Annals Entomological Society of America [Vol. IV, 

knowledge of specimens, the types of these descriptions grow 
in importance as the sum of our knowledge of species increases. 
The best description is not perfect, but, more often than not, 
deficient in some important taxonomic character. Hence the 
need of later systematists to refer to the type as the absolute 
standard of comparison. A nomenclature of types has accord- 
ingly been developed in recent years which is given the same 
importance as that which taxonomists attach to species nomen- 
clature. While less diversified than the latter, it should become 
of equal interest to the taxonomist, as it remains for him to 
apply it. 

With the close of the year 190G we have a series of five pri- 
mary types and four supplementary types designed to meet the 
needs of both systematist and type custodian. Some of these 
designations will possibly be disregarded or even found insuffi- 
cient; this depends upon the individual, whether he be "splitter" 
or "lumper." 

The first step toward a logical nomenclature of types was 
made when taxonomists began to set aside one of a series of 
specimens as the type proper, and to name the remaining 
specimens cotypes. Too often it had been found that a series 
which the protologist defined as one species actually represented 
two or more species. Hence the advisability of naming only 
one specimen the type and the others differently. The name 
"cotype", although used so universally, is in such case a mis- 
nomer and was finally set aside for the more pertinent and 
exact "paratype" — to signify specimens of the original series 
other than the type specimen. As the word "type" is subject 
to many interpretations according to the combination in which 
it is used, Schuchert in 1897 devised the word "holotype" — 
meaning "sole type" — for the single specimen on which a des- 
cription should be based. The name "cotype", however, was 
not discarded; its applicability only was limited. "Cotype", 
in its present interpretation, is properly applicable only in 
paleontology; for instance, when we have a fossil and its 
reverse. Another instance, from zoology, would be the follow- 
ing: two flies caught in coitu and not separated in death. If 
mounted together neither male not female can be called holo- 
type; there is no necessity of singling out one of the specimens, 
as there can be no doubt of the two belonging together. 

The following is a summary of type nomenclature: 

1911] Composition of Taxonomic Papers 207 

A. Primary Types. 

1. Holotype (H. T.) — A single specimen, or one selected of a series. 

2. Allotype (A. T.) — ^A single specimen of the sex not designated by 

the holotype. 

3. Cotype (S. T.) — Specimens of the original series when there is no 

holotype ( = syntype). 

4. Paratype (P. T.) — Specimens of original series when there is a 


5. Morphotype (M. T.) — A single specimen of the second form- 

described of a dimorphic sex. 

6. Lectotype (L. T.) — A cotype chosen after publication as holotype. 

7. Chirotype (X. T.) — Specimen on which a manuscript name is based. 

B. Supplementary Types. 

1. Plesiotype (P. t.) — Material on which subsequent descriptions or 

figures are based ( = apotype and hypotype). 

2. Neotype (N. t.) — ^A specimen from the same locality as the original 

type described or figured when the original type is lost. 

3. Heautotype (H. t.) — Specimen identified by the nomenclator or 

used by him for illustration, but not belonging to original series 
( = autotype). 

4. Plastotype (p. t.) — Plastic reproductions from type specimens. 

These must be casts. Models not included. 

The five prior names (1, 3, 4, 6, 7) for primary types are 
sufficiently "^simple and certainly not cumbersome for the sys- 
tematist. Yet it apjDears to me that one condition quite as 
important as the holotype has been overlooked; also a second 
one, which, if not general, still applies to certain orders of 

The first of these is easily apparent. Very many descrip- 
tions are based on one sex alone; often several decades pass 
before the unknown sex is discovered and described. Since 
this description is of primary interest to taxonomists, the speci- 
men on which this description is based in my estimation also 
merits a type name; and, what is more, should be classed among 
the primary types with the holotype. The second case is sex- 
dimorphism, common in a few orders of insects, rare in others, 
but still of such frequent occurrence that a type name for the 
dimorphic individual appears advisable. To designate these 
cases properly I have elsewhere (Bull. Milwaukee Museum, 
Vol. I, page 10, 1910) suggested the terms "allotype" — the 
other — -for the unknown sex, and "morphotype" — form — 
for the dimorphic form of a sex. 

208 Annals Entomological Society of America [Vol. IV, 

Allotype designates the sex not represented by the holotype. 
The allotype need not be described by the protologist (first 
describer) ; it can be contained in the original as well as in any 
subsequent description by other authors. Thus, if the protolog 
describes only a holotype male, the first female subsequently 
described is to be called the allotype; and vice versa. Mor- 
pliotype applies only to the second form' of a dimorphic sex. 
Here also the date when and the author by whom described are 
immaterial. (As the first form of a dimorphic sex will be repre- 
sented in the holotype or allotype, there may be some doubt as 
to the advisability of classing morphotypes among primary 
types. However, as both forms of a dimorphic sex are of equal 
importance to taxonomists I have placed morphotype in a posi- 
tion similar to the holotype and allotype.) 

Thus far few others than cataloguers have made use of the 
type-terminology here outlined. In fact, most of the terms 
were originated by them, since the thorough acquaintance with 
their subject gained by the compilation of catalogues has made 
them more susceptible to the various needs of taxonomy. As 
all of these terms are broad and permit of great latitude in inter- 
pretation and application, the systematist ought not hesitate 
to apply them. Past laxity in the treatment of types, and also 
in their preservation, has resulted in infinite confusion and has 
helped to increase synonymy beyond all reasonable bounds, 
so that in some orders the synonyms average 1.5 to each valid 

IV. Standards for Keys (Tables) of Genera and Species. 

1 . c? with appendage to hind tibia 2 

cf without appendage 4 

2. 9 with abdomen tufted 3 

9 with abdomen untufted Kilimanjaro 

3. Vein 6 usually curved in cf , 9 variable Popocatepetl 

Vein 6 usually straight Aconcagua 

4. 9 with abdomen untufted 5 

9 with abdomen tufted Matterhom 

5. Vein 6 curved Elias 

Vein 6 curved at end in cf Everest 

I defy anybody to reduce a specimen to its proper genus 
with a key of the foregoing type. Unfortunately, only too 
many of that sort exist and new ones are continually fashioned. 

A genus is the primary condition of taxonomy, and the use 
of secondary sexual characters for generic definition is an out- 
rage; an offense, which should not be condoned. Some of the 

1911] , Composition of Taxonomic Papers 209 

best taxonomists have placed their work in a questionable 
light by means of unsatisfactory tables like that given above. 
The only recourse in such cases is the original description, 
which is by no means such a simple proceeding as would appear 
on the face of it, as it often means a long, tedious search through 
many volumes. 

One may call the aid of the extended generic description, but 
the purpose of the key is to summarize what differences exist 
between genera. Tables are meant to be short-cuts through 
taxonomy; but I might as well try to run an engine on a rail- 
way which has one track alternately on each side of the ties, 
as determine a specimen from many generic tables. The use 
of geographical names in the key above is pertinent. It is 
just as difficult to climb those mountains as to determine speci- 
mens from some keys. Tables of the style outlined cause loss 
of tiftie, besides loss of temper. We are all human; and a 
scientist is not always the "dry, imperturbable fossil" the 
joke-antiquarians would have us believe. 

Among species tables we see many of similar nature. Yet 
here vagueness is excusable, while for an unsatisfactory genus 
table no valid excuses can be made. If the relations between 
two genera become too intimate, if distinctions fail — then the 
genera merge. 

Sexual characters are often the only ones that can be reliably 
applied in specific keys, and their use will be questioned by no 
one familiar with the difficulties of specific determination. 
Errors are possible everywhere, but they are offset by good 
work in other parts of the paper. Most often they result from 
a misconception of the specific value of certain characters. 
The aim, however, to compile a table of practical value will be 
easily apparent. 

Many of the difficulties of specific keys could be obviated 
by more care in the explanation of the essential characters 
used, their individuality, their variation, and their relation to 
others. But is there an excuse for the use of such terms as 
"larger species," "smaller species," "more slender," "more 
robust," and the like, in tables without in any way defining the 
limits of the terms? It is with feelings diametrically opposed 
to pleasure that I plod through a table of, say, 25 species, along 
lines indicated by "larger species," and "smaller species." 
What does the author mean thereby, I wonder? At which 

210 Annals Entomological Society of America [Vo.l IV, 

size does he draw the hne? My specimen is of moderate size 
and might be referred to either group. Therefore, is bulk the 
author's criterion? Or is length? Or width? Or odor? 

There are plenty of good, workable tables that will serve as 
models. An ideal table that would permit of "hard and fast" 
lines of division for species is, of course, impossible. But much 
could be done toward improvement by the elimination of indefi- 
nite terms from specific tables and sexual characters from generic 

V. Standards for Indices. 

Indices are the bane of scientific works. While their purpose 
is to facilitate reference to, and study of the contents of a vol- 
ume, it is rarely, indeed, that they achieve their purpose, 
because of their general insufficiency. Beginning with ordinary 
check-lists, bibliographies, travels, monographs, etc, taxonomic 
works are most often poorl}^ equipped as regards indices. 

It is impossible for any man to know all the species and 
genera of the average order. It is a fact, however, that just 
those publications which are greatest in volume and importance 
(taxonomical, ethological and otherwise) are the most poorly 
indexed. Some authors cite only genera in the index. Others 
feel that such method is insufficient and append the names of 
the species under the genera. While that is an improvement, it 
offers little aid to the student not familiar with the particular 

In this age of books, when it is possible to distinguish genera, 
species, synonyms, etc., each bj'^ various styles, sizes and impres- 
sions of types, the antiquated system of indices, as above re- 
ferred to, seems inconceivable. The trouble lies — so it seems 
to me — in the fact that authors seem to confound the index 
with a table of contents. 

To quote, "an index is a pilot through strange seas of 
thought. A book without an index is like a ship without a 
rudder. ' ' Continuing the simile — a book of entomology with 
generic index only is like an ocean-steamer with a canoe-rudder; 
and an index with the species names under the genera is like a 
ship with the rudder at its side. 

I need hardly assert that it is those books which are freely 
and carefully indexed that are most referred to. I feel much 
as the gentleman who said to me : "A scientific writer who does 
not care to make a complete and usable index to his works, 

1911] Composition of Taxonomic Papers 211 

should be prevented from writing at all! At the bottom of 
every insufficient index is not carelessness, but downright 

To set the standard for indices is not very difficult ; but the 
standard varies with the contents of books and papers. Here 
is the criterion : Since the aim of an index is to make the con- 
tents of a volume accessible to the reader, it should be so con- 
structed that it will permit access to the greatest possible 
number of references in the least possible time. In other words 
an index is a medium of saving time. Hence an index should 
not be merely a carelessly jumbled summary of the contents, 
but a carefully arranged alphabetic list of all names, facts and 
captions in the volume. This includes technical as well as 
popular names, generic as well as specific names. 

There is such a thing as over-indexing. The author must 
use his judgment as to the amount of detail he desires to index. 
Also, unnecessary repetition should be avoided. One fact, how- 
ever, is patent; that if the author wishes to see his work con- 
sidered at all as a work of reference, he must supply it with a 
good index. I, for one, do not care to use poorly indexed books, 
and consult such as rarely as possible. To say the truth, 
I consider it a personal affront, when upon purchasing a book, 
I find myself maltreated to several hundred pages of facts and 
names, and a two-page index. The author h"as no cause to 
treat his readers as if their brains were ware-houses; that they 
need but read his book and file away the contents together with 
the exact page number, etc., for future reference. By pur- 
chasing and reading a book I am doing the author a twofold 
service. And if I remember some of the statements and quote 
the book as an authority, the acme of the author's expectations 
is then reached. More he has no right to demand. But a 
starved index is inimical to progress, since few men will care 
to quote when they are unable to find the passages from an 
insufficient index. 

When is an index desirable? One friend has stated this 
succinctly: "Any taxonomic paper citing more than fifty 
names should have an index of its own. " This seems reasonable 
to me. An index of fifty names, run in two columns, eight point 
on a ten point base, would occupy less than the ordinary four 
by seven page of our journals. Because of the practice of 
societies and institutions to send reprints to an author for pri- 

212 Annals Entomological Society oj America [Vol. IV, 

vate distribution, this special index seems more than justified; 
unless the author expects his associates to supply the index 
privately. But this is expecting too much. Take, for instance, 
some of our well-known entomologists, who receive hundreds of 
reprints in a year, among them contributions exceeding 100 
pages. It is astonishing, how few of these larger papers are 
supplied with an index at all; at that, the indices are mostly 
of the Spartan type. Should these men undertake the neces- 
sary clerical work and compile the missing indices? True, many 
of these men keep card-indices of their specialties. But what 
of workers on more than one branch of entomology, or zoology? 
To keep card-catalogues — hence general indices — of their wide- 
ly distributed interests would necessitate the employment of a 
clerk throughout the year. 

I close with the classic from Pope, "He who knows how to 
prepare a good index, holds the eel of science by the tail. ' ' 

VI. Standards for Titles. 

In logical order the title should have been treated first. 
But since the title is usually the last thing written by an author 
for his contribution, so let its place be among the last in the 
order of standards. 

Take any entomological journal in hand and glance over the 
titles of papers. Many of these will sound much like the fol- 
lowing examples: "A Revision of the Genus Popocatepetl; 
Some New Species of Orizaba; A New Aconcagua; A New 
Variation and the Life History of Kilimanjaro alta; etc." 
Occasionally one meets a title like the following: "A New Genus 
and Species of the Family Sierra"; and indeed a rarity is 
"New Species of the Order Andes." 

In North America alone there are about 70,000 described 
species of insects, distributed in approximately 8,000 genera 
(probably more). Nevertheless, everyone is, as a matter of 
course, expected to know immediately from the lucid "Genus 
Popocatepetl" just where the genus belongs, to what family, 
to which order. Everyone is expected to be familiar with all 
of the 8,000 genera and to have no difficulty at all in placing the 
genus revised or enlarged, as indicated by the title. And even 
considering that there are about 500,000 specific and 80,000 
generic names in zoology, "Popocatepetl" is too important 
not to be as well known as "pater" and "mater." 

1911] Composition of Taxonomic Papers 213 

Especially in taxonomic entomology the saying holds good: 
"Familiarity breeds contempt" — for others. Some taxonomists 
appear to become so obsessed with their particular specialty 
that other orders or families of insects do not exist for tfiem. 
There are 18 other orders after Comstock, 30 others after 
Handlirsch (restricted to Pterygogenea — winged insects) ; yet 
these are of little importance beyond the fact that they exist 
and that some foolish people bother about them. So taxo- 
nomists of a certain type would have us believe. We are lucky, 
indeed, if with indignant compassion they will cite the family 
in which the order occurs; indignant, because "those bar- 
barians" do not happen to take any special interest in their 
particular branch. 

Let us go a step farther. There are eighty-two families in 
the order Coleoptera, sixty-one in Diptera, about seventy-five 
in Lepidoptera, about seventy in Hymenoptera, not to speak of 
Hemiptera, Neuroplera, Pseudoneuroptera, and other orders. 
A conservative estimate would show over four hundred families 
of insects in North America alone, distributed in nineteen 
(Comstock) or thirty-one (Handlirsch) orders. Most of these 
families average three to four subfamilies to each family, and 
two tribes to each subfamily. Figuring on this basis there are 
1200 subfamilies and 2400 tribes of insects. And this for North 
American insects only! What of the orders, the families, the 
subfamilies, the tribes, the genera, of fishes, of mollusks, of 
birds, of mammals, of crustaceans, etc. in North America? 
What of their number in the entire world ? Not all our articles 
are confined to a single fauna. The Central and South Ameri- 
can faunas are beginning to be explored more thoroughly, as 
shown by the ever increasing number of articles upon the 
regions named. 

And yet, on an average but six out of twenty titles cite the 
family, and but one of twenty the order. Of course, the fact that 
the journal is specially devoted to entomology, gives me a clue 
to the position of the genus ; accordingly I know that the paper 
is an entomological paper, but that is all. But what of journals 
dealing with natural history in general ? How can I know from 
the title whether the genus belongs to botany or to zoology or 
paleonotology, whether it is a paper on insects or canaries, on 
mollusks or angle-worms? 

An hour spent in a scientific library in the classification of 
articles would be an educative influence for aU those who neglect 

214 Annals Entomological Society of America [Vol. IV, 

the mention of family and order in their articles, The difficul- 
ties they would meet — such as antiquated catalogues, under- 
indexed catalogues, or, as in some cases, the entire lack of 
catalogues — would forever cure them of this apparently trifling 
but nevertheless momentous negligence. Even when there are 
good catalogues at hand, it is a complex proposition to place a 
genus. For the terminology of some orders, such as Diptera, 
and Hymenoptera, Coleoptera and Hemiptera, etc., is, in part, 
alike; the necessary consultation of both text and catalogue 
in such cases causes an irksome and avoidable loss of time. 

The solution of all troubles is so simple, so obvious — in fact, 
it is inherent in the subject — that it seems strange why taxo- 
nomists have not adopted the simple means. But one ento- 
mologist is known to me who in all of his papers inserts the 
order name in his titles. That is the solution : Insert the order 
of the insect, bird, mammal, or whatever-it-be behind the genus 
and family name in the title. This holds good also for mor- 
phological, ethological and other papers as well as for those 
dealing solely with taxonomy. 

VII. Standards for Reprints. 

This chapter does not properly belong in this consideration. 
But since reprints form an important part of the specialists' 
literature, a few words on the topic may be of interest. 

Sometimes I receive reprints of articles published by "Enig- 
ma" University; that is a tangible fact. The paging of the 
reprint is the same as originally published; that is another 
tangible fact. But I look in vain from page to page in the 
endeavor to discover the number or j^ear of the volume, the 
month of publication, etc. That editor who arranged the 
reprint of an article sent me, published in nineteen-something 
on pages 260-290 of a certain periodical, yet paged the separate 
1-30; and carefully efl^aced all reference to the name of the pub- 
lication, the year or number of the volume, the year and month 
of publication; — that editor, I say, deserves no honorary men- 
tion. After guessing at the probable publications in which the 
article might have appeared, I looked over the recent volumes 
of many and ultimately succeeded in ^finding the exact place, 
page and time of publication. I owe that editor thanks, since 
through him I was led to other articles of high interest; but I 
spent an entire evening in trying to find out "What's which" 
in the reprint. To be full}^ consistent, the editor should have 
effaced the title of the article itself. 

1911] Composition of Taxonomic Papers 215 

To be sure, this was an extreme case. Yet that in these 
"enlightened" days, after years of discussions, protests and 
recommendations, there should be men who retain the benighted 
idea that it is preferable to change the paging of reprints from 
the original — this seems hardly conceivable. Why the change 
at all? No advantage is gained thereby. On the contrary, 
it is a disadvantage for workers who are not constantly in touch 
with all the leading centers of scientific work and who have no 
large scientific library at their elbows. For these it results in 
tedious correspondence, and this most often when there is 
little time to be spared for these irksome labors. 

One lucid individual went to another extreme. The travels 
of a certain explorer, together with the scientific results of his 
collections, as monographed by various specialists, were pub- 
lished in a large scientific journal. As all of these contributions 
were finally to be collected in a separate volume, and as the 
paging of this volume would be just as important for reference 
as that of the journal, the editor thought of a "happy" solu- 
tion of all difficulties. Namely, the original paging of the con- 
tribution as it appeared in the journal was retained for the 
reprint; the future paging of the volume was also put in; and 
to meet all contingencies the reprint was given a special paging 
of 1 — 50 or other. Unfortunately, this genius forgot to note 
which was which, so that, as the printer's folio number and the 
publisher's file number are at the bottom of each page besides 
the three numbers above, I now have my choice between five 
numbers for page reference. 

As a rule reprints do not suffer from surplus information 
as in the preceding case ; they usually lack part of the necessary 
information. This lack in most cases is the absence of the vol- 
ume number (or the year of the volume) from the reprint, or 
the year of publication, or both. Sometimes the two are given, 
but the name of the publication is nowhere indicated. The 
benign opinion that every scientific worker is familiar with the 
size of the volumes, the style of composition and the issues of 
"the four-hundred" leading scientific publications, — this opin- 
ion is, of course, founded on long experience and hence must be 
considered sound. If I receive a reprint that contains the 
year and number of the volume, but not the title of the publica- 
tion itself, it is, therefore, a simple proposition to locate the 
correct journal from the size of the page and the style of com- 

211) Annals Entomological Society of America [Vol. IV, 

position, as there are only about three hundred others among 
the "four hundred" that resemble it. 

Often the title of the journal is present and the number of 
the volume given, but not the year of the volume. The latter 
is omitted because it is a matter of common knowledge that the 
institution or society began its journal way back in the forties 
and that a new series is begun with each score of years ; so that 
the tale, "Reprinted from the Enigmatical Journal, Series 4, 
Volume 17" will tell me all that is necessary to be told. From 
the number of the volume I ought to infer the year of the volume 
and if I am too much of an "ignoramus" as not to know such a 
monumental fact as the year a certain society or institution was 
founded, — well, then "look it up!" 

Similarly, if I read 1906 on a reprint just received, I am to 
know intuitively that that means the year of the volume, not 
the year of publication; that the contribution had been in the 
hands of the editor since 1905, but owing to the press of legis- 
lative matters on the state printer could not be published until 

It appears ridiculous that a matter intrinsically so simple, 
and extrinsically of such vital importance as the correct mark- 
ing of reprints should be so carelessly treated. Or is there really 
a living editor who would consider the puny additional (?) 
expense of the line on the reprint giving all the needed informa- 
tion? Penny wise, pound foolish. Can a simpler solution be 
found than "Reprinted from the Ecstatic Journal, Series 6, 
Volume 14, pages 28-67, 1910 (Publ. May, 1910)"? 


The scope of inatters that are left to our imagination, divini- 
tion and intuition by scientific papers is monumental. A cata- 
logue of merchandise that does not describe the ware and state 
its prices would be flung aside instantly. Yet for science any- 
thing, no matter how poorly constructed, how poorly presented, 
should be acceptable. Science should lead the world. But if 
science in general cannot apply more logic to its methods than 
taxonomists apply to taxonomy, its leadership will be short- 
lived. This may be a harsh and pessimistic view ; but I believe 
that I do not stand alone in this attitude. 

Again referring to the merchandise simile — imagine to your- 
self a catalogue of merchandise, say furniture, that would not 
bear the proper legend on the cover; further, that the pages 

1911] Composition of Taxonomic Papers 217 

•contained nothing else but names of furniture — no illustrations 
of the same, no measurements, no prices quoted; — imagine the 
action of the man receiving it! Certainly no other place than 
the paper-basket would be accorded it. And certainly many of 
the articles of our journals are little better as far as usable 
information is concerned than the furniture catalogue just 
referred to. Is it with reverence that we remember such names 
as Smith and Walker of British Museum fame? And yet some 
systematists appear to have chosen them as patrons and models 
for imitation. They succeed only too well in imitating them, 
and occasionally outdistance them. 

One may say, these are all minor matters. That is true. 
But their aggregate forms an imposing array. One drop in a 
■cup will not make it acrid; but a number of drops will change 
it into a cup of bitterness. So with entomology. One little 
•carelessness does not amount to much; but many will fill even 
the most ardent student with feeling akin to disgust. 

Science is no longer in its infancy and we have a right to 
demand advanced methods of work. The desire for improvement 
is innate to all men. I have never heard of a writer (at least 
in science) who was well satisfied with what he had written. 
Literary critics say, "An author is his favorite reader"; but 
self-satisfaction is short-lived, more so in science than else- 
where. Hence the attitude of scientific workers toward their 
work may be defined as "a minimum of self-conceit with a max- 
imum of scruples. " Writers do not confess these qualms of the 
intellectual conscience to the public, but reserve them for some 
private interchange of confidences. Unfortunately, the ratio 
■of these qualms decreases, not inversely, but in the same ratio 
that the system and methodical effort of the worker decreases; 
so that the most conscientious workers are usually most diffi- 
dent as regards their own work (all the more, as those contri- 
butions requiring the greatest amount of labor and time gen- 
erally show the least for it) , while the careless workers have few 
misgivings of their efforts. I have an inkling that some day to 
come a contribution will have to be passed upon by a commis- 
sion of scientists (like so many examination papers) before they 
are declared acceptable to science. 

Cooperation and centralization (to a certain extent) are 
desirable. There ought, in fact, to be a scientific clearing house 
somewhere in this beautiful world, and I hope that it will be 
achieved some day. 


The Annals of the Entomological Society of America will be 
published by the Society quarterly and will include the Proceed- 
ings of the Annual meetings and such papers as may be selected 
by the Editorial Board. 

Papers may be submitted to any member of the Editorial 
Board and should be as nearly as possible in the form desired as 
final, preferably typewritten, and illustrations must be finished 
complete ready for reproduction. Plates must not exceed 5x7 
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must be original, complete and previously unpublished and, ex- 
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to publish preliminary announcements or notes. Authors will 
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Morgan, Anna H — Mayflies of Fall Creek 93 

TowNSEND, C. H — Announcements of Further Results 

Secured in the Study of Muscoid Flies 127 

HiNE, J. S. — Robberflies of the Genera Promachns and 

Proctacanthus 153 

Burgess, A. F. — Locomotion of the Larva of Calosoma 

Sycophanta 173 

Webster, R. L. — Notes on the Pear Slug iSi 

Severin, H. H. p. and H. C. — The Mechanism in the 

Hatching of the Walking Stick, Diapheromera 

femorata Say '. 1S7 

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Gove:-n Entomological Publications _ . . - . 192 

Muttkowski, R. A. — The Composition of Taxonomic 

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Volume IV. Number 3. 



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Eolered a< second class matter April 1 1, 1908, at the Post Office at Columbus, Ohio, 
under the Act of Congress of March 3, 1679. 

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Davis, John J. — Studies on Aphididae II 20 

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the Larva of Corydalis Comuta '. 25 

Nelson, Jas. A. — Evolution and .Adaption in the Palpus of Male Spiders 16 

Webster, F. M. — Investigations of Toxoptera Graminum and Its Parasites . . .25 

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Biological Building, O. S. U., Colombos, Ohio. 



The Entomological Society of America 

Volume IV SEPTEMBER, 1911 Number 3 


By William A. Hilton. 

Concerning the insects, many extensive works have been pub- 
Hshed upon the nervous system from early times down to quite 
recently. The work of Dujardin, '.50, may be said to be a 
starting point. Numerous papers by Villanes from '87 to '9.3 
give general accounts of the structure, but nothing very def- 
inite as to the distribution of individual nerve termination and 
origin within the ganglia. The extensive work by Saint-Remy, 
'90, is also a somewhat fragmentary account of numerous forms 
of tracheate head ganglia. Other earlier papers dealing with 
cephalic ganglia in particular are those of Newton, '79, and 
Packard, 'SO, and in more recent times we have the valuable 
works of Kenyon, '96, and Haller, '04. In connection 
with the structure and relationships of abdominal ganglia, the 
investigations of Binet, '94, and Benedicenti, '95, should be 
mentioned ; and for a summary of the form and structure of the 
insect nervous system, the general work of Berlese, '97, is 

Although there are numerous and extensive papers dealing 
with the structure of insects, very few give a very complete 
account of the whole nervous system of a single species and 
practically no single work treats of the larval centers in much 
detail, although numerous papers take up the development and 
some as Bauer, '04, consider the transformations of larval into 
the adult conditions. 

The external anatomy and general distribution of ganglia 
and nerves of Corydalis have been studied by Krauss, '84, and 
by Hammar, '08. The relations of the trachea to the nervous 
system and their distribution within it by Hilton, '09. The 


220 Annals Entomological Society of America [Vol. IV, 

present paper is a continuation of the study of the nervous 
system in the larval form and, although not as complete as 
might be wished, it is at least a start in the direction of a clearer 
comprehension of the insect central nervous system, under- 
taken for the purpose of preparing for a studj- of the finer 
structure of the nerve cells, and for experiments upon their 
metabolism and function. 

The methods employed were various. For obtaining the 
best idea of the general distribution of nerve cells and fibers, and 
the tracts of which they are parts, intra vitam methylene blue 
injections were used. Beautiful results w-ere obtained at 
times, but it was only after hundreds of specimens were gone 
over that much was learned as to the organization of the gan- 
glia. Sectioning methods with the usual fixers and stains gave 
fair results and the methods of Golgi and Cajal were tried, also 
those of Villanes and Kenyon. All of these gave good prep- 
arations except the Golgi method which I hope to try again at 
another time. There were difficulties in the way of fixing and 
staining because the ganglia are inclosed in chitin and because 
of the numerous tracheal vessels, and in the larger ones it was 
not possible to get perfect whole mounts. Sketches were 
made from the methjdene blue preparations both before and 
after fixation and in the first stages of the work peripheral 
nerves were traced by means of gross dissections. 


The abdominal ganglia, eight in number are quite uniform 
in appearance and general structure with the exception of the 
eighth or most caudal. The first abdominal is separated by only 
short connectives from the third thoracic, and the seventh is 
even closer to the eighth. The seven first abdominal ganglia 
have quite uniformly on each side, two large nerve trunks 
connected with them, a cephalic lateral and a ventral more 
caudal branch. The eighth ganglion has four pairs of branches 
leading into it from the caudal end of the animal. 

Specimens were injected with methylene blue and nerves 
traced to the periphery and from here followed into the ganglia 
as nerve tracts as far as possible. In an earlier study on th-.^ 
nervous system of larval insects I found that in some cases 
some of the more cephalic branches connected with the ganglia 
were in large part if not totally sensory, that is arising from 

1911] Nervous System of Corydalis 221 

bipolar nerve cells and nerve plexuses, from tactile hairs and 
from the surface of the hypodermis. In Corydalis at various 
times during several years I have tried to determine the motor 
and sensory parts of each peripheral trunk for the purpose of 
following them into the central nervous system. To some 
degree methylene blue stain is of a differential value in deter- 
mining the nature of nerve trunks, for very often the first 
neurons to take the stain are sensory, while motor fibers and 
cells are often slower to turn blue. But this method is not 
absolutely sure, for there is great variability in the staining 
reactions of different individuals. The only sure way of telling 
whether a given branch is motor or sensory is by tracing the 
nerves to their endings in muscle fibers or from their origin in 
bipolar sense cells at the periphery. The tracing of a motor or 
a sensory nerve or tract is not possible in a large number of 
cases because the stain is incomplete or too dense, but 
occasional selectively stained preparations enable one to make 
positive if not complete statements in regard to nerve trunks; 
that is to say, one can determine surely from a specimen that 
a large number of branches of a certain nerve are all motor or 
all sensory, but it would be impossible to say with perfect 
assurance that the nerve was pure motor or pure sensory 
because some fine terminations might remain uncolored, 
especially in the case of a stain which was good for sensory 
terminations, for there would be a strong probability that some, 
at least of the fine motor ends would not show. 

The work of Hammar, 'OS, on the nervous system of Cory- 
dalis has been very helpful, and the general description of the 
nervous system given by him is so complete that I shall not need 
to spend time on the gross anatomy of the various ganglia, and 
in speaking of the several branches of the ganglia 1 shall follow 
his terminology. 

There are three chief branches breaking from the Lateral 
trunk of each of the first seven abdominal ganglia, their method 
of branching from this trunk and from each other is somewhat 
variable, but these three main parts are easily recognized. 
Branch 2 is large and comes off quite near the base of the lateral 
trunk, runs caudally a short distance and then disappears 
between muscle fibers in a ventral direction. I could not 
determine it to be anything but a motor branch although some 
of the fibers from it are among the first to stain and some of 
them pass not into the ganglion connected with the nerve 

222 Annals Entomological Society of America [Vol. IV, 

trunk, but run directly up to the next ganglion by the way of 
the connectives, in a tract which from its other connections in 
other species and in this form, and from its staining reactions, 
I took to be sensory. Branch 3 is long, it runs up to the dorsal 
side of the animal and is without doubt mixed motor and 
sensory, containing fibers which supply dorsal muscles anti 
fibers which come from the hypodermis. Branch 4 runs into 
the lateral appendage and seems to be sensory, for the most 
part at least. Besides these, there are two minute branches, 

1 and 5, running out to the trachea, according to Hammar, 'OS. 

The ventral trunk runs caudally and ventrally, branches 1, 

2 and 3 run to more and more caudal portions of the ventral 
side of the animal and seem to be entirely sensory, branch 3 
runs to some extent also into the lateral appendage, while 
branch 4 runs into the tracheal gill and was the only one traced 
into it. So this whole ventral trunk seems to be for the most 
part sensor}'. 

The eighth abdominal ganglion seems to be made up of at 
least two centers fused, there are four main trunks entering it 
on each side below and all of these so far as could be determined 
are both motor and sensory. Trunk (a) is most lateral, (b) 
a ventral trunk corresponding to the ventral one of other ab- 
dominal ganglia, (d) a more median one supplying lower dorsal 
and ventral portions of the body and (c) median, with a large 
branch which runs back up the intestine. 


(Fig. 5.) 
The nerve cells of the periphery have already been figured 
in an earlier article, Hilton '02. The functional cells of the 
ganglia both thoracic and abdominal appear to be much of the 
same type in methylene blue preparations, uni- or bipolar nerve 
cells, one of the processes or branches of which may run out 
quite a long distance before they break up into a number of 
terminations, the other portion usually breaks lip into branches 
near the cell body. Indications of multipolar cells were seen 
in some specimens but with these usually all of the processes 
but one were verj' small and hard to trace very far. In addi- 
tion to the functional neurones of both large and small size, 
there were in all of the ganglia, numerous neuroblasts, or smal- 
ler cells \\dth slight protoplasm about the nucleus, and neurog- 
lia networks. 


Nervous System of Corydalis 



(Figs. 1 and 2.) 

By means of methylene blue preparations it was possible in 
some more deeply stained specimens to trace the main tracts of 
fibers within the ganglia and within the connectives and in 
lighter stained specimens the distribution of special tracts 
and even individual fibers. At times the cells stained as well 
as the fibers at other times only fibers were colored. 

Fig. 1. Figure of the 7th and 8th abdominal ganglia from methylene blue 
preparation. Dorsal side. A few nerve cells are shown in black. The chief nerve 
trunks show with their fibers. The central "Punktsubstanz" of the ganglia dotted. 
Some of the larger tracheal tubes shown as thick solid black lines. 

The caudal end is down in this and the following figures. x.30. 

Fig. 2. Sixth abdominal ganglion from ventral side. Methylene blue. x30. 

Stained or unstained, the central region of each ganglion is 
more opaque or darker, due to the nerve fibers crossing and 
terminating in this region. This forms on each side a central 
body made up of two oval masses more or less fused into one at 
the middle line, the "Punktsubstanz" of some authors. The 

224 Annals Entomological Society of America [Vol. IV, 

nerve fibers of the connectives when stained in a mass form 
deep Hnes apparently running straight through the center of 
the ganglia, these longitudinal bundles of nerve fibers seem to 
be a little broader before entering and after leaving the central 
mass. The nerve trunks in deeply stained specimens send 
masses of fibers into the ganglia and in the case of most of the 
fibers, the region where they seem to terminate is in the central 
part of each ganglion. This is true of all the ventral fibers 
and of most of those from the lateral trunk, but a few of the 
latter, and some fibers from the second branch of the lateral, 
run up into the edge of the ganglion only, and then straight up 
the connective to the next ganglion above. In the case of the 
eighth adbominal the four nerve trunks enter the fibrous central 
mass from below, those most medially placed seem to be con- 
tinued up through to the connectives and to be largely contin- 
uous with them in deeply stained specimens, while the more 
lateral trunks are lost sight of as they enter the central portion 
of the ganglion, although some of the fibers from the more 
laterall}^ placed nerve trunks pass through the edge of the 
ganglion without communication with its cells and pass up 
the outer side of the connectives on either side to the next 
ganglion above. There are then two masses of fibers entering 
each center but the last, those of the connectives and those of 
the nerve trunks. I will first take up those of the connectives. 
Beginning with the seventh abdominal ganglion great masses 
of fibers enter, and it is possible to distinguish ; (a ) Fibers which 
run straight through without terminating. There seem to be 
great numbers of these, but this is due in part to the fact that 
when fibers do terminate in a ganglion they end at various 
levels. These fibers can however individually in a number of 
cases be traced through a ganglion without endings of any sort 
within it, just how far some of these may run without termina- 
tion is a question, but there was no difficulty in tracing them 
through three ganglia and there is no reason to doubt that 
they may be longer than this. Those most easily followed 
were usually of larger size than the rest, (b) Fibers from below, 
terminating within the ganglion. Of these there are several 
sorts: (1) Those ending in the lower part of the "punktsub- 
stanz" on the same side. (2) Those ending on the same side 
above. (3) Those crossing over towards the opposite side 
from below. (4) Those crossing over to the opposite side above. 

1911] Nervous System of Corydalis 225 

In those entering from below some run straight in and end 
in the caudal region of the central fibrous mass, while in many 
.specimens fibers from the outer side of the connectives sweep 
sharply in towards the center of the lower part of the ganglion 
to end near the middle line, either on the same side or just 
over it. (c) Fibers from above. In general there are similar 
bundles of fibers to those traced from below: (1) Those end- 
ing in the lower part of the ganglion on the same side. (2) Those 
ending on the same side but in the cephalic portion of the 
ganglion. (3) Those crossing to end in the lower part. (4) 
Those crossing to end in the upper part. 

In the case of fibers ending in the ganglion from the ceph- 
alic direction, none were seen forming such a dense sweep into 
each center from the sides of the connectives, although there 
were a few fine ones of this sort. Most of the fibers leave the 
" punktsubstanz " to run in the connectives without great devia- 
tion from a straight course, (d) Fibers passing into the con- 
nectives from cells within the ganglion. There may be dis- 
tinguished in many of the preparations cells with their fibers 
well stained, the more central of these may be more clearly 
seen in some cases. Some of the larger more central cells seem 
to be merely for association within the ganglion, with all of 
their processes ending within it. Others send one main pro- 
cess up one connective and another down into one of the other 
great masses of fibers. Other cells of medium or small size, 
located chiefly at the sides of the ganglion send one long process 
into one of the nerve trunks while the other shorter process 
may run for a short distance in the connective trunk or be lost 
in the central mass of the ganglion. 


These have already been spoken of to some degree. Most 
fibers of both cephalic and ventral nerves seem to enter the 
central part of the ganglion and are lost track of in the "Punkt- 
substanz," but both the lateral and ventral trunk-fibers are 
continued into the connectives in the cephalic direction at 
least, and possibly to some extent in the caudal, although this 
was not determined. In the case of the lateral trunks of all 
the abdominal ganglia, there is a possible sensory tract enter- 
ing the cephalic edge of the nerve center without coming to the 
central "punktsubstanz" or having any communication with 

226 Annals Entomological Society of America [Vol. IV, 

nerve fibers, running along the outer side of the connective and 
for the most part ending in the basal portion of the ganglion next 
above, near or across the middle line. A similar tract to this 
has been described coming from the two most lateral trunks 
jn the last ganglion. 

These fibers which enter from cephalic lateral trunks seem 
to stain among the first and in the case of some other insects 
were found to come from bipolar sensory cells at the periphery, 
and I still think that they are to some extent sensory, but these 
tracts which have no communication with the cells of the gan- 
glia with which they are connected are not all of the sensory 
fibers of each nerve center, for the ventral branches have many 
sensory fibers and these do not follow exactly the same path, 
and in the case of the first seven abdominal ganglia many of 
the fibers could be traced from the branch 2, which so far as 
could be determined was a decidedly motor trunk. 

Fibers other than those coming from cells on the opposite 
side to run into the branches as motor axones, are directly 
supplied by cells on the same side, long branches from certain 
cells run into the various motor trunks while the other termina- 
tions are in the "punktsubstanz." 

Fibers from the periphery or from sensory cells enter the 
ganglion from both main trunks and are of the following 
groups": (a) Those ending within the ganglion to which the 
trunks are connected, the exact termination of these T could not 
make out, but some at least ended near the central part of the 
ganglion, although very often arborizations of the terminations 
could be traced both on the same side and on the opposite side. 
Fibers entering straight from below in the last abdominal broke 
up into branches near the middle line with arborizations in the 
central margin of the ganglion. 

(b) Those passing from one ganglion to the next without 
sending branches to the center to which the nerve trunks are 
connected, some of these fibers may run past one or more 
ganglion, but the most of them form a definite tract from the 
peripher}^ by way of lateral trunks, running on the outside of 
the connectives, and turning sharply in towards the middle 
line in the caudal portion of the central mass of fibers, to end 
here or a little higher up, or to cross over and end in the "punkt- 
substanz" of the opposite side not far from the middle line. 

1911] Nervous System of Corydalis Til 

(c) Those passing from the periphery into the nerve trunks 
and having extensive arborizations in the ganglia to which they 
are connected and then passing on to another ganglion with 
arborizations in it. Only a few of such fibers were distinguished 
one in connection with the Sth ganglion was the clearest case. 
A nerve fiber from tho periphery was easily traced into the 1st. 
lateral trunk, a branch from this fiber was given off" in the ceph- 
alic and lateral region of the ganglion, this fiber could be traced 
into the " punktsubstanz " of the nerve center, some of its arbori- 
zations ending on the same side and one branch was traced to 
the cephalic region of the other side, while the main fibers passed 
up the connective and ended by arborizations in the "punkt- 
substanz" of the ganghon next above chiefly on the same side 
in the caudal region. 


Individual cells and fibers were not so easily traced by this 
method, but general masses of fibers and the location of cell 
groups were determined. 

All of the ganglia, connectives and nerve trunks are inclosed 
in a chitinous envelope which in many cases is very close to the 
nervous tissue but usually separated by neuroglia cells. This 
envelope is especially thick about the connectives just before 
and just after they enter a ganglion, it appears as a uniform 
mass in section with large and smaller openings where trachea 
penetrate it. 

In places under the chitin of the ganglia, especially on the 
dorsal side, there are large spaces with little or nothing in them 
but delicate neuroglia networks. The trachea radiating in the 
chitin covering the connectives and ganglia have already been 
referred to; as stated in a previous paper large branches and 
fine tracheoles run to the nervous system and are distributed 
to all centers and their branches. These are superficial or run 
in the chitinous sheath, and the deep, supplied in part by the 
superficial twigs but chiefly by larger special branches and 
enter the ganglion and connectives. In these connectives it 
is easy to see numerous openings, large and minute between the 
masses of nerve fibers, and in cross section the air tubes are 
shown to be fully as numerous as one would expect from a 
study of surface views where all the trachea were made to show. 
Tracheal tubes within the ganglia are particularly noticeable 

228 A^inals Entomological Society of America [Vol. IV, 

in the centers of bundles of fibers and most easily seen in these 
traced from the connectives. The exact place and method of 
termination was not determined. Injections of fluids into the 
ganglia by way of trachea failed to penetrate any of the finer 

All of the abdominal ganglia seem to be of practically the 
same type, but individual variations occur. 

In all of the nerve centers the cells are grouped for the most 
part on ventral and lateral portions of the ganglion and towards 
the caudal end, a few cells occur on the dorsal side especially 
near the middle line and these are often quite large. 

Description of 4th abdominal ganglion traced by sections 
beginning at the caudal end: 

The connectives entering from the ventral side are easily 
followed as distinct longitudinal masses of fibers well up into 
the ganglion, these connectives as well as others in other parts 
of the nervous system are composed of numerous closely 
packed longitudinal fibers, scattered between these are the 
openings of trachea, when the ganglion is reached the chitin 
for each of the connectives becomes fused into one mass and 
farther in the central portion of chitin between them disap- 
pears and the two bundles of fibers are more or less crowded 
against each other. Farther up into the ganglion the fiber 
bundles do not occupy all of the area under the chitin because 
large spaces on all sides occur and then soon cells in a single 
layer are found close to the wall of the ventral side, and then on 
the dorsal side a very large cell is found wedged in between the 
two bundles of fibers. Some of the cells of the ventral side 
may be seen at this level sending fibers into the two longitudi- 
nal bundles. The single layer of cells on the ventral side 
becomes a double row of medium and small, and the large cell 
of the dorsal side gives way to a group of small ones and there 
comes to be on the ventral side two groups of fibers running 
more transversely, probably made up in part from fibers con- 
nected with the cells appearing on the ventral side. 

Farther up these ventral nerve cells extend out laterally so 
that numbers of them might be seen from the dorsal side. No 
cells are left for a distance on the mid-ventral line, and they 
disappear from the mid-dorsal line also to some extent, but 
before they are gone fibers can be traced about the connective 
bundles and to the cell regiqn of the ventral side. At this 

1911] Nervous System of Corydalis 229 

level there are nerve fibers seen between the cells on the ventro- 
lateral margins of the ganglion and fibers connected with these 
regions of the nerve center join the bundle from the cells on the 
dorsal side, on the ventral median side of the ganglion, while a 
third runs in from these cells into the central part of the longi- 
tudinal fibers. We have then at this level three transverse 
bundles of fibers crossing from the lateral cell groups, a dorsal, 
a ventral and median and a little farther along we have also a 
bundle of fibers running across the section but from the dorsal 
to the ventral side and uniting to some degree with the three 
right and left commissures. Other little branches from these 
main ones and other tracts from the lateral cell groups also 
invade the longitudinal bands from the connectives. 

A little above this level again on the ventral side a single 
layer of cells appears in the middle line and no cells are seen on 
the dorsal side except laterally. 

A little above this, the large ventral trachea enter passing 
through the cell layer and breaking up into numerous branches. 
The central fibrous mass of the ganglion is largely made up of 
longitudinal strands in all levels so far and besides the com- 
missures mentioned there are usually a number of fibers crossing 
irregularly both dorso-ventrally, laterally and obliquely espe- 
cially at about this last level. None of them are large and the 
great mass of fibers remains longitudinal. It is at about this 
level that the ventral nerve trunks come off from the lateral 
and ventral sides of the ganglion from the central part of the 
latero-ventral cell mass, just before the tracheal trunks are 
reached. Fibers from this trunk may mingle with the cells of 
this region and are also continued into the central mass of 
fibers of the ganglion. 

Beyond this point the cells become thin again especially 
ventrally and also laterally, the central thickest part of the 
ganglion is now reached and the fibers form a rather large dense 
mass. Longitudinal ones may still be seen mixed in with 
numerous lateral and transverse strands all bound up together 
into a dense fibrous mass with no very marked special tracts or 
strands except for quite a well marked short broad median 
commissure of fibers connecting more intimately the two 
already well fused masses of each lateral half of "punktsubstanz." 

Slightly beyond this, the cells have about disappeared, only 
a few remaining at the dorso-lateral edges of the ganglion. 

230 Annals Entomological Society of America [Vol. IV,. 

Beyond this something of the central commissure remains, many 
of the other crossed fibers in the central part of the ganglion 
have disappeared. A bundle of fibers partly transverse and 
partly fused with the central longitudinal bands begins to be 
seen on either side of the ganglion ventrally, these are partly 
mixed with the main longitudinal tracts. They are endings of 
the bundles of the lateral nerves to be followed later and might 
be called lateral nerve tracts. At this level a few scattering 
cells on the ventral side and two small dorso-lateral groups, one 
on each side of the ganglion indicate about all of the cell masses 
seen lower down, while in the mid-dorsal line a new group of 
dorsal cells makes its appearance and sends fibers through the 
central part of the ganglion as a central tract which breaks up 
laterally and can be traced to various parts of the central fiber 
mass of the ganglion. For several sections these fibers become 
quite prominent and the central commissure seems to be lack- 
ing, then as this central tract disappears higher up, another and 
a better marked commissure comes to view running trans- 
versely through the center of the ganglion from side to side. 
At this level cells again come into view laterally. The ventral 
tracts of the lateral nerves become more prominent and there 
is a dorsal band of fibers close to the edge of the "punktsub- 
stanz " on the dorsal side. This last is parallel with the median.. 

Slightly beyond this a few cells are seen on the ventral side 
laterally, two of the same commissures, a dorsal and a median 
may be seen, but the lateral cells have disappeared to give place 
to the entrance of the fibers of the large lateral nerves. These 
fibers for the most part run directly into the lateral nerve tract 
noted above when it was seen more caudally. Beyond this and 
beyond the entrance of the lateral nerve, a few cells are seen 
laterally, one or so in the mid-dorsal line, and the dorsal and 
median connectives disappear and only a few tangled fibers 
replace them, although for a few sections the great sweep of 
transverse fibers is continued from side to side, from the lateral 
nerve tract. 

Above this no commissure or cross fiber of any sort connects 
the lateral halves of the ganglion and a small group of nerve 
cells comes to lie on the middle line and dorsal and ventral to it. 
At the line of separation of the lateral halves, the tracts of the 
lateral nerves can be distinguished as a dense mass on either 

1911] Nervous System of Corydalis 231 

side of the longitudinal fibers which are continued out into the 

Above this as the cells disappear and we come clearly into 
the region where there are only longitudinal tracts, these may 
be followed and they are indistinguishable from other fibers of 
the connectives. The reason why the lateral tracts could be 
told from the longitudinal for such a distance was because they 
seemed denser and stained more deeply. The fibers in the 
cephalic connectives have about the same arrangement as the 
caudal ones. 

In other abdominal ganglia, ventral and lateral groups of 
nerve cells were more clearly seen contributing to the com- 
missures and the central tracts. Some of the fibers of the 
lateral trunks end in the central portion of the ganglion, prob- 
ably in cells. 

The tract of the lateral trunk needs a word of additional 
comment. In preparations made bj^ a method that removes 
the cells and all but the denser fibers so that little more than a 
skeleton of the fibrous framework is left, it is found that a 
transverse portion connecting the two sides of the ganglion is 
much denser than other parts of the fibrous mass and under the 
highest powers of the microscope, this seems to be very finely 
granular as well as fibrous and is continuous from side to side 
between the nerve trunks. This same fine granular substance 
with fibrils in it was traced up into the connectives a short dis- 
tance, and as many fibers are seen to end in this region it may 
be due to a dense grouping of their endings that there is a 
deeper color at such a place. Similar substances to this only 
in more isolated portions is found in other parts of the ganglion 
and in other nerve centers. In specimens stained with ordinary 
hematoxylin there is no differentiation between this substance 
and the general fibrillar mass. 

The eighth abdominal ganglion is similar to the others 
except that the connective fibers begin within the ganglion and 
there are more commissures developed. The first lateral 
branch can be easily traced out into the connective on the out- 
side, fibers also deeper in go on up the connective, while still 
others enter the ganglion and are distributed to all parts of one 
side and probably also across to some extent, as there are 
numerous cross connections, by means of at least three of four 
well marked commissures, besides irregular fibers. Other 

232 Annals Entomological Society of America [Vol. IV, 

branches also send fibers to the central mass, some of these run 
straight through, while others seem to cross in commissures 
or end. 

In general then, there are in each abdominal ganglion, cells 
on the ventral caudal region, on the lateral sides, and a few on 
the median dorsal side. These cells surround a central fibrous 
mass made up of strands running longitudinally through the 
ganglion from the connectives and best marked in the cephalic 
and caudal parts; fibers running across from side to side, these 
run in about three commissures, a dorsal, a ventral and a 
median and at various cephalic and caudal levels these com- 
missures are interrupted. The lateral nerve trunks may be 
seen to contribute largely to the formation of the large ventral 
commissure. The other cross connections seem to be more 
exclusively from cells on the sides of the ganglia and from these 
cells also other cross or diagonal fibers may be followed. 

The dorsal group of cells which seems to be to a large 
degree for association, sends fibers through the ganglion to the 
cells of the lateral and ventral groups, so that these fiber tracts 
may be found above or below the commissures penetrating to 
the opposite side, or part way through when the median com- 
missure is present. 


Methylene blue method. (Fig. 3). 

The three thoracic ganglia are quite a little larger than the 
abdominal and the branches come off differently. 

There are on each side three main trunks the most cephalic 
of these has its most cephalic branches pure sensory, but No. 2 
was not determined, also No. 1 of trunk B or the middle trunk 
seems sensory while other branches of the middle trunk are 
more or less mixed and the last which goes into the leg is also 
mixed. So then the more cephalic nerves are sensory while the 
rest seem to be mixed. The exact nature of the two parts of 
the last or leg branch was not determined, but there was no 
reason from the staining reactions to indicate that they were 
of greatly different composition. 

In the thoracic region as in the abdominal, the main trunks 
easily took up the stain, but here greater difficulty was encoun- 
tered in surface studies because of the larger opaque mass of the 
ganglion. Cells and fibers were however made out and found 


Nervous System of Cor'ydalis 


to be in a general way similar to the conditions found more 
caudally. The main tracts of the connectives and of the nerve 
trunks enter the central portion of each center as in the abdom- 
inal region, but their distribution within was harder to make 
out. There were tracts entering the last thoracic ganglion 
from below, leaving it again as in the abdominal centers. 

Fig. 4. 

Fig. 3. Third thoracic ganglion from below. Methylene blue. x30. 

Fig. 4. Connective branch leading off between the 2d and 3d thoracic ganglia, 
nerve fibers from above and below enter the nerve trunk from the connective. 
Also large and small nerve fibers shown. Methylene blue. x45. 

Tracts from the first abdominal pass up the outside of the con- 
nective and cross over into the middle line, but from the third 
thoracic to the second, and from the connectives of the second 
to the first no such tract was clearly recognized. Fibers enter- 
ing laterally both from motor and sensory nerves all pass in 
towards the central part of the ganglion. In other words there 
was no indication of a tract passing from cephalic branches into 
the edge of the ganglion to run without termination up the out- 
side of the connective to the next center. But there was an 
indication of fibers passing through or into one ganglion from 
the one below it. 

In the cephalic part of the thoracic ganglia fibers coming 
from above may some of them be traced as a fine tract ending 

234 Annals Entomological Society of America [Vol. IV, 

in the cephalic portion of the ganglion. Other than these 
differences, there were no essential ones between these nerve 
centers and those of the abdominal region. 

In regard to the arrangement of cells as shown by methylene 
blue, it was found that the lower ventral and lateral regions 
had the greatest number, great masses of them, with many 
more cells than in the smaller ganglia. For the most part 
similar arrangements of individual ribers were seen. Nerve 
cells sending fibers directly into motor trunks, cells of medium 
or rather small size, were observed, but these were few in num- 
ber. Most of the cells seen had their processes running into 
the "punktsubstanz" of the ganglion. Large and smaller asso- 
ciation cells were found as in the lower regions and of various 
sorts such as already described for them, some at the surface of 
the ganglion other at the edges of the "punktsubstanz." 

Between the third and second and the second and first 
thoracic ganglion, there are branches off from the connectives, 
a pair between each of these, and between the subesophageal 
and the first thoracic there are two pairs. The upper of these 
last were not so well stained in any of the preparations but all 
of the others were quite well colored and found to be motor. 
These branches when studied as to their composition did not 
differ much from each other and in each one, fibers could be 
seen descending to run out the nerve trunk from the ganglion 
next above and also from the ganglion below. These two 
tracts of fibers entering the lateral trunks were clear and dis- 
tinct from each other for quite a distance into the nerve trunk, 
(Fig. 4). 


(Plate XV, Figs. 1-4.) 

The internal structure of the thoracic ganglia is much more 
complicated than the abdominal, due to the fact that the larger 
branches from the more numerous nerve cells are more inti- 
mately woven together, and it was practically impossible to 
follow commissures or tracts very far except in a very general 
way. However, a general description as detailed as seems 
necessary will be given of one of the thoracic ganglia, the first. 

From above the connectives which enter as in the abdom- 
inal ganglia are in every way similar. Not many cells are seen 
scattered in the upper part of the ganglion, then two large 

1911] Nervous System of Corydalis 235 

groups appear one on eaci'; side laterally and a small ventral 
group. (Fig. 1-3, Plate I). These masses at the sides of 
both large and small cells are at least three deep. The three 
groups a little farther along become united by a single row of 
cells which farther up becomes double layered and all the cell 
groups are not distinguishable in the single mass. There are 
also at about this level as a part of this mass a few cells in the 
mid-ventral line between the bundles of fibres of the connectives. 

Farther up, the connective tracts are less clearly all longi- 
tudinal fibers and the lateral part of the nerve cell mass gives 
way for the entrance of the first or most cephalic of the three 
nerve trunks, the fibers of which pass into and mingle as trans- 
verse and dorso-ventral fibers in the connective tracts. The 
fibers of this nerve are very extensive and may be followed into 
the center of the ganglion, both dorsally and ventrally. Fibers 
from the ventral cells on either side of the ganglion enter the 
center of each lateral half from below and are there lost and 
partly pass into the nerve trunk. Fibers from the cells in the 
mid-ventral line, which cells form a wedge shaped mass at 
higher levels between the connective masses, run to the dorsal 
side of each of these masses of longitudinal fibers, and from 
here circle about to become associated with the fibers of the 
nerve trunks and with other more median strands on each side 
of the ganglion and with the strands described above which 
come from the ventral mass. Slightly beyond this part and 
nearer the center of the ganglion the two central masses of 
fibers or connective masses become fused together, the cells 
disappear and commissures, a dorsal, a ventral and a median, 
connect to some degree the sweeps of fibers already described. 
(Fig. 4, Plate XV, just above this level.) 

Farther down, two commissures, a median and a dorsal are 
seen but numerous fibers cross the middle line at many levels 
and angles. Farther on but one commissure can be noted, 
a ventral, but many other fibers cross at different angles and 
the whole lateral portion of the ganglion is a dense system of 
complicated interlacing fibers having a dense meshwork. On 
the lateral part of each ventral half the fibers stain darker, 
probably due to more numerous fine branches in this region 
and on the dorsal median line a little wedge shaped group of 
cells makes its appearance, the only cells of this region. These 
send their fibers through the center of the ganglion to the ven- 

236 Annals Entomological Society of America [Vol. IV, 

tral side, while a central commissure crosses these to end in the 
tangled mass of fibers on either side of the ganglion. Farther 
along, these dorso-ventral bands a little one side of the middle 
line do not cross the now larger central commissure, but run in 
to it as do the other fibers from the ventral side, running 
from the more deeply stained ventral mass already spoken of. 

Farther along and at the level of the next nerve, three com- 
missures, a ventral, a dorsal and a median may be again recog- 
nized while the fibers of the middle nerve both end in the lateral 
portions of the fibrous mass and contribute to the three commis- 
sures. In this level only a few scattering nerve cells were seen. 
Beyond this a ventral, almost a lateral group appears again on 
each side and fibers from these form a little arch about the now 
smaller mass of darker staining fibers. On the mid-dorsal line 
fibers from this arch and others from these cells also ramify 
into all parts of the ventral portion of the ganglion. Along from 
this the dorsal part comes to be separated into two separate 
masses of longitudinal fibers of the connectives again. Far- 
ther along the arch becomes in its dorsal portion fused into a 
median commissure which soon disappears as the cleft between 
the connectives becomes deeper and reaches way down to the 
now small area of deeply staining substance which now forms 
a ventral commissure. The ventral cell group has become more 
lateral at this level and another large group has come in just 
dorsal to it, but still only on the side. In the mid- ventral line 
also, there has come in a small new group of cells. 

The last nerve trunk comes to be associated with this com- 
missure of deeply staining fibers on the ventral side and farther 
along fibers also pass freely into it from the lateral group of 
cells which has been spoken of as coming in more dorsally, this 
for a time remains distinct from the other more ventral groups. 

Along farther these cell groups unite to form a large thick 
single lateral mass and from them more fibers run into the com- 
missure of deeply staining fibers and " Punktsubstanz. " 

Soon after this the commissure breaks through as the two 
connective bundles separate, each with a little of the darkened 
mass which soon disappears as do the cells of the ganglion. 

Although the above description is only a very general one, 
it will be seen that the ganglion is more complicated than the 
abdominal, but the general plan of arrangement and structure 
is as in the abdominal region. The nerve cells as in the abdomi- 


Nervous System of Corydalis 


nal ganglia are chiefly grouped in the caudal, cephalic and 
ventral regions and may be seen to take direct part in the for- 
mation of commissures as well as diagonal strands. While 
dorsal cells on the median line and ventral median cells, send 
fibers through the ganglion dorso-ventrally, as well as associa- 
tion fibers to different tracts and lateral groups. In both 
thoracic and abdominal ganglia dark staining masses made up 
of very minute fibers fused together are chiefly found on the 
ventral side and associated with a ventral commissure. 


Fig- 5- 

Fig. 5. Nerve cells from the central nervous system, (a) Motor nerve cell 
from the 3d thoracic ganglion, (b, c and d) Association cells from the. same, 
(e) Cells from the brain. xlOO. 


Annals Entomological Society of America [Vol. IV, 


(Figs. 6 and 7, Plate XVI, Fig. 5.) 
This ganglion is larger than the others described, and is less 
flattened and less easy to study from the surface. The branches 
have already been traced quite well to the periphery and I will 
only mention them briefly. 

Fig, 6. Fig. 7. 

Fig. 6. Subesophageal ganglion from the ventral side. Methylene blue. x30. 

c connective with brain. 

CO commissure 

mx maxillary 

m mandibular 

1 labial 

g gustatory 

Fig. 7. Subesophageal from the dorsal side. x30. 

The caudal portion of the ganglion becomes thick soon after 
the connectives have entered. The cephalic lateral portion of 
the ganglion is connected with the supraesophageal above by 
two large connectives, but smaller than those from the 1st 
thoracic ganglion. These cephalic connectives or crura cere- 
bri are connected together a short distance away from the gan- 
glion by a cross branch or commissure. 

1911] Nervous System of Corydalis 239 

From the cephalic end there are three pairs of large nerve 
trunks, the mandibular, the maxillary and the labial. The 
mandibular is the largest the labial the smallest and most 
ventral. All appear to be mixed nerves, both sensory and motor. 

Either side of the middle line on the cephalic border are 
two small nerves, the gustatory, which are motor in part at least. 
On either side of the ganglion not far from its central portion 
is a small ventral nerve and not far from the connectives near 
the entrance of the caudal tracheal tubes are the small salivary 
nerves. I know nothing of the composition of these two last 

The dense central mass of the ganglion prevents one from 
tracing nerve fibers very deeply in surface preparations, but a 
few more fortunate specimens gave now and then a fiber or a 
tract which could be easily followed. In general with the nerve 
trunks and connectives of other ganglia, these bundles of fibers 
entered the central portions and like them, too, the nerve cells 
were chiefly grouped on the sides with scattering cells on the 
dorsal and a denser mass on the ventral and caudal portions, 
but in this the dorsal side has more cells than was usual with the 
other ganglia. The same arrangement of cells and fibers was 
noticed as in others, that is, most of the peripheral cells could be 
seen to send their processes into the central portion. Fibers 
from the connectives above and below could be traced through 
the ganglion, but there were such masses of them that it was 
difficult to tell whether they were branched or not. 

Fibers from the lower connectives were seen to end in the 
caudal portion of the " punktsubstanz " : (a) On the same side, 
(b) Crossing over the middle line. These were both superficial 
fibers and resembled those in the bases of the abdominal gan- 
glia. Probably deeper fibers end higher up. 

Fibers running down the upper connectives run : (a) Down the 
connective to end in the central portion of the ganglion; (b) 
Down the connective to end in the caudal region of the ganglion. 

Probably among both of these groups of fibers there are some 
which cross over into the opposite side of the ganglion. 

Fibers running down the connectives and crossing over to the 
opposite side through the commissure connecting the crura cere- 
bri: (a) Cross over in the commissure to the opposite side and 
run down to end in the upper or lower portions of the ganglion. 

240 Annals Entomological Society of America [Vol. IV, 

Two other sorts may be given although no complete fibers were 
traced through such a course; 

(b) It seems probable from the specimens that fibers cross to 
the opposite side in the commissure and run over to the oppo- 
site side of the ganglion: 

(c) Probably some fibers cross in the commissure and run 
back to the brain. 

Fibers running straight through the ganglion from above 
and from below were not traced but it is very possible that such 
are present as in other ganglia. 


The mandibular branch sends its fibers into the cephalic 
dorsal border of the " punktsubstanz." Some of its fibers seem to 
end here, others pass in deeper. 

The maxillary sends its fibers into the very center of the 
upper half of the ganglion and here some of them seem to end or 
cannot be traced farther in surface views. This is true of the 
more cephalic branch of the maxillary in part at least, while the 
rest of. the fibers of this and those of the caudal branch are 
traced in laterally a little farther down. 

The fibers of the labial nerve; some of them run in deeply 
about where the branch enters the ganglion, others go down 
farther and may be traced as far as the place where those of the 
ventral nerve trunk enter the mid-lateral portion of the central 
fibrous mass. 

The salivary nerve fibers run in and can be traced to near 
the point where the ventral nerves were. 

The small gustatory nerves run some distance down into the 
ganglion from the point where they take their exit and a motor 
nerve cell was found sending out its axon directly into this 


The connectives which run up to the subesophageal ganglion 
are much like the others described. As the caudal portion of 
the ganglion is reached these two longitudinal tracts of fibers 
become fused although they may be distinguished from each 
other. A group of nerve cells appears on the lateral sides, and 
a group of large ones on the median side dorsally, some of these 

1911] Nervous System of Corydalis 241 

penetrate in between the tracts and a few cells appear ven- 
trally on the median line, while the cells become more numerous 
laterally. The salivary nerves enter latero-ventrally and unite 
with the mass of longitudinal fibers. The cells disappear dor- 
sally, but some are between the mass of fibers of each con- 
nective and the lateral cell group has become more ventral. 
There is at this level a transverse commissure on the dorsal 
side and fibers running down ventrally in the ventral line. 
Farther along the fibers do not so many of them run from 
dorsal to the ventral side and a median commissure comes to be 
formed. More cells come in laterally and ventrally and these 
sending their fibers into the central mass contribute to its 
complexity. These cells also run into the large but ill-defined 
median commissure. 

Two little spots of darker more dense fibers come in on the 
ventral side and fibers from the ventral cells form an arch 
about them. 

Fa,rther cephalad the median commissure becomes less well 
defined. The dorsal is lost and a median group of cells comes 
in dorsally again. Fibers from the ventral and dorsal cells, 
especially the former go in curved sweeps to the dorsal and 
ventral sides of the fiber mass, fibers also run in laterally from 
the lateral cell groups. Farther along no clear commissure can 
be seen, but sweeps of fibers cross from both sides, those of 
opposite sides interdigitating to some degree. The ventral 
darker mass of fibers mentioned a short time ago has now 
become a transverse mass and is larger, being joined by fibers 
from the labial and ventral nerves. At the level of this entrance 
only a few scattering groups of cells are seen. 

The dark fiber mass becomes expanded to the center of the 
' ' Punktsubstanz. 

The broad cerebral cruri are reached. From the central to 
the dorsal side laterally four small groups of nerve cells mostly 
small, with now and then a large one are seen. Fibers from the 
cerebral crus can be traced to the center of the fiber mass and 
into one or more of the several irregular masses of dark fibers. 
Dorsally and ventrallj' fibers cross from side to side and run 
diagonally from the dorsal to the ventral side. Farther along 
a ventral commissure of dark fibers is present, some of its strands 
reaching up into the dark fibers in the direction of the com- 
missure and farther along breaking through it. A short dis- 

242 Annals Entomological Society of America [Vol. IV, 

tance cephalad of this point the ventral mass disappears and the 
lateral halves of the general mass of fibers becomes distinguish- 
able once more as the upper region of the ganglion is reached. 
At this upper region, lateral cells are no longer seen, there is 
however, a small group ventrally placed, either side of the 
middle line and a small dorso-median mass. Fibers sent in 
from these curve up to run into the crus which may also at this 
level be seen to receive fibers from the middle and opposite 
parts of the ganglion by the way of a group of fibers just one 
side of the middle line and a group running from the center of 
the lateral central mass. 

Just beyond this last level at the place where the maxillar}^ 
branch enters, a little group of nerve cells comes in between it 
and the crus. Fibers from this large maxillary 'nerve run into 
the crus, into the central and ventral portion of the ganglion and 
apparently across to the other side, while many of its fibers are 
lost in the deeper staining central masses. 

Farther along the ventral cells become much more abundant, 
a wedge shaped group G-7 layers thick with a few large cells. 
A few cells come in on the mid-dorsal line and some come in 
latero- ventrally just above where the maxillary nerve joins the 
ganglion, and some of these cells seem to contribute directly to 
the nerve. 

The large mandibular nerve joins the ganglion on its upper 
border, fibers come to it from ventral and dorsal sides of the 
ganglion and connections with the darker fiber masses in the 
center can be traced. Cells are now in masses both dorsally 
and ventrally as the cephalic end of the ganglion is approached 
and some of these at least seem to contribute directly to the 

The above description is a very general one, only the main 
features of structure and arrangement were spoken of. The 
complexity of the ganglion is such that a general summary of it 
follows : 

(a) Cells 

The cells at various levels dififer greatly. Beginning at the 
caudal end and passing forward there might be recognized 
about three main dorsal cell groups one after another which 
fuse and separate from each other at various levels. The median 
ventral cells are at first also separated from the other groups 

1911] Nervous System of Corydalis 243 

but farther up they grow out laterally to become continuous 
with the lateral and dorsal cells at various levels while they be- 
come absent from the mid-ventral line, then become united 
again on the cephalic region where all the cell groups are joined 
together. In intervals where these groups are not in distinct 
masses a few scattered cells are often found. 

(b) Connectives 

Ventral connectives. Fibers from these run straight into the 
ganglion for a short distance until the central, tangled mass is 
reached. Only a few of the fibers in the central part of the 
ganglion can be seen to take a straight course through it. 
Many others run straight or nearly so for a short distance and 
then turn off sharply to one side. Fibers from the connectives 
seem to end at all levels and in practically all parts of the 
central fibrous mass and to be contributed to by cell masses 
especially on the ventral side, but also clearly on the dorsal. 
These fibers coming in from both sides of the ganglion at differ- 
ent levels and as single fibers or groups add considerably to the 
complexity of the ganglion as does the fact that many of the 
fibers from the connectives which run through to the crura 
cerebri and nerves do not always take a straight course or run 
to the same parts of the nerve trunks. Sweeps of fibers for 
instance, can be traced quite straight up on the ventral side of 
the ganglion and then may be seen to turn over to the dorsal 

Crura cerebri. These have fibers from the caudal connect- 
ives but not nearly all from them can be traced into the crura, 
for they are smaller and have their own special fibers which 
come from almost every part of the ganglion. The cells in 
various parts seem to furnish many of these, some of which 
come from the same side, but single strands were followed 
running in the direction of the crura which were from the 
opposite side. Fibers may also be seen to sweep back into it, 
probably from the mandibular trunk. 

(c) Nerve trunks. 

Mandibular. Many of the fibers of this end in the first 
part of the fiber mass. A few apparently run into the crura. 
Some fibers could be traced from near the median central part 
■of the ganglion in a line with the lower connectives. Some 
•came to it from cephalic median cells. 

244 Annals Entomological Society of America [Vol. IV, 

Maxillary. Fibers were followed into this from the lower 
connectives and from the upper parts of the ganglion. From 
this nerve trunk some fibers seem to end near the junction of 
the nerve with the central "punkstubstanz." 

Labial. This is made up from fibers which enter the ven- 
tral central portions of the ganglion, just above the entrance of 
the small ventral nerves. They maj' be traced from the con- 
nectives up and from the upper portion of the ganglion down 
into these trunks and ventral caudal cells evidently contribute 
fibers to the mingled mass which is connected with these 

(d) Commissures. 

The commissures connecting the crura cerebri have fibers 
which cross from one side to the other in the case of descending 
or ascending strands. No other kinds were recognized although I 
think there is a strong probability that some fibers merely 
cross and do not descend at all. 

Within the ganglion there are a number of commissures 
connecting the lateral halves. Some of these are of straight 
fibers, others are closely woven deep staining masses. Dorsal, 
ventral or median commissures are found at almost every level, 
especially ventral ones, although not always clearly marked. 
A longitudinal section through the whole ganglion shows from 
three to four main commissures, a cephalic, a caudal and twO' 
median ones. 


(Figs. 8, 9, and PI. XVI, Figs. 1-4.) 

The brain is made up of two large ovoid masses distinctly 
marked from each other on the the middle line. It is connected 
on the ventral side to the subesophageal ganglion by means of 
the short, broad crura cerebri. All of the larger nerves come 
out laterally and of these there are three main trunks, the only 
ones to be considered at this time. 

Three portions of the brain may be made out each con- 
nected with these trunks. The most dorsal is the protocere- 
brum, and it is also the largest and best marked and connected 
with the optic nerves which divide on each side into seven, 
branches one for each ocellus. 


Nervous System of Corydalis 


The middle lobe of the brain or the deutocerebrum is the least 
marked of any and its nerve trunk the antennal, is the smallest 
of the three. It enervates muscles at the base of the antena as 
well as sense organs in it and so is mixed. This lobe is best 
seen on the cephalic and dorsal side and not at all on the cephalic 


Fig. 8. 

F'g- 9- 

Fig. 8. Supra- and sub-esophageal ganglia with their attached nerves and 
ganglia. The brain is turned over cephalad. The sub-esophageal ganglion is 
dorsal. x.30. 

a antennal fn frontal nerve 

c clypeolabral v vagus ganglion 

f frontal ganglion o optic 

Fig. 9. Cephalic view of one half of the brain. Methylene blue. x.30. 

246 Annals Entomological Society of America [Vol. IV, 

Finally the tritocerebnim is well marked as a little lobe just 
dorsal of the crus giving off the rather large clypeo-labial 
trunk which with the arched nerve comes off as one. From the 
distribution of this it seems probable that it is mixed. 

This arched nerve runs ventrally and cephalad to unite at 
the middle line with the one of the opposite side in the frontal 
nerve ganglion. A branch from this small nerve center runs for- 
ward as the frontal nerve, another runs back on the dorsal 
surface of the esophagus to the small vagus ganglion, which 
sends two branches farther down the alimentary canal. 

General form of the Brain from Methylene blue. 

In successfully stained preparations almost all parts of the 
brain, especially the parts in from the nerve trunks are seen to 
be covered with nerve cells, both large and small. Those just 
under the chitin seem to quite completely incase the central 
dark staining portions of the ganglion. This central dark mass 
in each well separated lateral half of the ganglion is roughly of 
the same general shape as the surface. In the main part out 
from the median portion there is a lobe deep in and opposite the 
ocular nerve, this is in the central portion of the ganglion and 
connected with it, but extending down into the tritocerebrum is 
another lobe almost as large near the crus. 

Partly separated from the central lobe of " punktsubstanz " is 
a spherical mass of dark staining substance and out from this a 
little distance in the ocular lobe and beyond its constriction 
from the main part of the protocerebrum is another little mass 
of deeply colored material. About each of these last little 
masses of " punktsubstanz " a special arrangement of cells is seen, 
while over the surface of the main portion of dark substance on 
every side the cells form a thick covering. 

Fibers running up the crura may be traced into the ganglion 
in its dorso-caudal region. Some apparently run only to the 
lower portion of the "punktsubstanz," others may be followed 
farther up and are lost in the central area. Fibers can also be 
traced to the central portion of the ganglion, to the medial 
portions and probably freely ramify all through the central 

Near the middle line of the ganglion some large cells on the 
surface were found with long processes extending down long 
distances in the direction of the crura and probably were con- 
tinued into it. 

1911] Nervous System of Corydalis 247 

The great bulk of fibers connected with the crura seem to 
take origin or terminate in the central portion of the "punkt- 

The ocular nerves enter the protocerebrum through large 
nerve trunks which form a decided lobe on the surface of the 
brain. At the junction of this ocular lobe with the ganglion 
there is a little area of deeply staining substance mentioned, 
before and back of this are nerve cells, and also a few cells on 
the eye side of the mass. These may be seen to send their 
processes into a dark mass and in towards the main part of the 
brain. Fibers run out the nerve from the ocular lobes' deep 
staining mass and into the spherical body before mentioned and 
into parts of the " punktsubstanz " near it, the former are pro- 
cesses from cells located near the ocular "punktsubstanz." 
About the spherical mass may be seen many nerve cells whose 
fibers are connected with it. 

Nerve cells on all surfaces of the protocerebrum are very 
numerous and may be seen sending their processes into the 
central fibrous mass of the ganglion. 

The deutocerebrum is less marked than the other two 
neuromeres and the fibers of its nerve, the anteniial, come in 
closer to the clypeolabial segment of the brain than the ocular 
portion. The fibers of the antennal nerve can be traced as a 
distinct band for nearly one-half of the distance from its 
entrance to the middle line, where they seem to end in a mass of 
deep staining fibers of the clypeolabral trunk where it joins 
the main central portion and here at least some fibers can be 
seen to end well towards the caudo-ventral portion of the 

The tritocerebrum is best marked in the dorso-cephalic side 
of the ganglion where it lies over the crus. The fibers of its 
nerve seem to be of two sorts. The labral part is often stained 
while the arched nerve portion is clear. Both branches enter 
the ganglion and plunge at once into the mass of deeply stain- 
ing fibers. Not quite so many cells were stained overlying this 
region in the specimens prepared. Some of these sent fibers 
more or less directly into the central mass while others as in 
other surfaces of the brain seemed to be association cells in a 
small area. 

248 Annals Entomological Society of America [Vol. IV, 


Connected with the arched nerves somewhat cephalad of the 
brain is the frontal ganglion. This in well stained preparations 
may be seen to have a central deep staining mass surrounded 
by nerve cells, the processes of some of which run into the 
central mass, while those of others run out from the cell, and 
down into the nerve which runs under the brain and connects 
this with the smaller so-called vagus ganglion. This last is 
like the former only smaller and fewer cells surround the central 
mass, some of the fibers run from this and probably also rather 
directly from the cells of the ganglion, down and out the two 
caudal branches. Occasionally the two lateral ganglia of the 
esophagus take the stain but their connections or structure was 
not especially studied. They seemed to differ from the other 
two ganglia, as they showed from the surface no nerve cells, the 
whole body taking on a uniform deep blue color. Sections 
showed them composed of very many cells closely massed 

Sections of the frontal ganglion show a small mass of cells 
quite well filling the caudal end, a few larger, but mostly 
smaller cells of the same general sort found in other places. 
Of these there were about two large and eight smaller ones at a 
level where the ventral nerves come off on each side of the 
"Punktsubstanz," although farther cephalad than the place 
where these nerves are seen from the surface. Farther cephalad 
where there are only three or four large cells, fibers cross in 
various directions in the central portion of the ganglion. A 
large cell for instance was seen to send a process into the center 
of the ganglion where it broke up into a number of branches. 
Fibers cross in the various directions but most run longitudi- 
nally. At a level where a branch to the frontal ganglion arises, 
there are no nerve cells, the central part of the ganglion is 
divided into three masses of longitudinal fibers by trachea and 
cross fibers. This division is continued only for a short 

Farther along a dorsal and a few small ventral cells come in. 
The central mass of fibers is rather uniform, but made up of 
both cross and longitudinal strands. A little farther cephalad 
three cells come in dorsally at about the level where the arched 
nerves come off. Fibers running from side to side connect 

1911] Nervous System of Corydalis 249 

these nerves through the center of the ganglion. 

The so-called vagus ganglion has a central mass of fibers and 
at its central part a nearly double row of cells closely packed 
about this central fibrous mass, these cells are continued down 
from the center a short distance, especially on the dorsal side. 


(Plate XVI, Figs. 1-4.) 

Only the main features of the structure of this complicated 
organ will be given at this time. Many of the elements of the 
brain of the adult may be present in the larva but for the 
proper interpretation of these it will be necessary to follow up 
this work with studies on the ganglia of pupae and adults. 

As in the other centers, a central fibrous mass forms the 
bulk of the organ and about this central " Punktsubstanz " 
nerve cells are grouped in great numbers on practically all sides 
but the ventral. As in the other ganglia, large and small nerve 
cells and small neuroblasts are found, the latter are especially 
abundant and occur in great masses. Besides these, filling in 
between and in places where there are no nerve cells is the 
neuroglia network, which is often continued to the layer of sur- 
face supporting cells just under the chitinous sheath of the 

In sections we may recognize the dense staining parts seen in 
surface views to be masses of fibers more or less complexly 
arranged in the central and ventral portion, more or less par- 
alleled by straight bands of the entering nerve trunks. Some 
of the special denser masses of fibers have already been described 
from surface views as that in the ocular lobe just as it joins the 
brain and in from the little spherical area just within and 
beyond this point. These two masses although quite separate 
from each other dorsally, ventrally and laterally are centrally 
connected by fibers and are also connected to each other to a 
less degree in the same way. The central fibers described in 
connection with the crura cerebri, are continued down into the 
labral lobe, but the deepest mass is in the central portion 
dorsally where it is somewhat lobed because of groups of cells 
on the surface and due to the processes of some of these cells 
passing down into the center. In this central "Punktsub- 
stanz" either side of the middle line, is a well marked denser 
group of fibers, a rod of substance projecting from those on the 

250 Annals Entomological Society of America [Vol. IV, 

dorsal side of the brain. This runs down to the mid-ventral 
line and there branches into two parts, one ventral, the other 
dorso-lateral. Farther along each of these parts run caudally 
as a single piece, one ventrally, the other dorso-laterally and 
the middle portion of the rod and afterwards the dorsal part 
disappears in section because of the curve in it. Later the 
lateral part disappears and the mid-ventral portion extends in 
towards the middle line to meet, but not unite with its fellow of 
the opposite side, running caudally in this way some distance, 
just above a ventral fibrous commissure and below a broad 
central one. In other words this body is a long slightly curved 
rod standing up in the ganglion with its base divided into two 
portions of which the lateral is shorter, the median longer 
and extends in towards the middle line. These represent the 
stalks and roots of the "mushroom bodies," the cup, such as 
described by Kenyon and others is not present and the special 
cells if developed were not recognized. This stalk and root of 
fibrous substance seemed to have a lighter core, that is in sec- 
tions it gave to some extent the appearance of a tube. The 
fibers which compose it are very densely massed together. 
Preparations in which the tissues were allowed to macerate 
showed them to be little aflfected. 

On the cephalic margin of the brain as on the dorsal side, 
the central fibrous mass as a whole is lobed as already spoken of 
and masses of cells fill in over these. The cell groups are 
difficult to describe in detail. The whole dorsal and lateral 
portions of the ganglia are covered with them, both large and 
small and in places many cells deep. There are almost no cells 
on the ventral side of the brain. 

Beginning laterally and dorsally we have about the spherical 
mass of fibers back from the ocular lobes, masses of cells, on 
the dorsal, ventral and mesal sides. A peculiar condition of some 
of the dorsal and ventral sides of this mass is the appraent epi- 
thelial character of some of the cells. Most of these are very 
small and are probably neuroblasts. The epithelial character 
is especially marked in two places on each side because there 
are little cavities one dorsal and one ventral in the outer por- 
tion of the circular mass of fibers. (Plate XV, Figs. 1, 2, 3.) 
There are some fibers from the two lateral groups of cells just 
described which run both to the mesal group and out the ocu- 
lar nerve. This is also continued dorsally and forms all along 

1911] Nervous System of Corydalis 251 

the dorsal part of the ganglion a thick layer, in the middle 
region of the brain. 

A group of large cells occurs dorsally either side of the mid- 
dle line, most of these point ventrally or centrally and send fibers 
to the crura cerebri, to the commissures and to the central por- 
tions of the fibrous mass; theirs are the longest fibers recognized 
from any cells in the brain. 

Out laterally and ventral to the ocular lobes in the region 
of the antennal lobe and just above the crura is a small group of 
nerve cells, sending fibers into the crura and into the fibrous 
substance near that region of the brain. 

In the cephalic region the cells surrounding the spherical 
mass may be seen divided into a dorsal, a ventral and a median 
group of small cells, already mentioned, while larger ones fill in 
on the dorsal side and are part of the general dorsal mass. 
These and the median masses run together and separate again 
at various levels, groups of smaller and larger cells often alter- 
nating, and these are continued on the cephalic and caudal sides 
of the ganglion. One of the most marked is a small group of 
cells surrounding a curved lobe of the central mass of the 
ganglion and continuous with cells on the cephalic side of the 

Fiber Tracts in the Brain. 

(1) The labial. Fibers seem to end chiefly in the dense 
fibrous mass located in the labial lobe. A few fibers could be 
traced doubtfully into a dorso-lateral group of cells. 

(2) Antennal nerve. Fibers from this end in cell groups 
either side of it. Fibers pass down ventrally into the lateral 
central part of the ' ' punksubstanz ' ' in large masses where some of 
them end, others cross to the ventral side and run in strands 
back in the main tract of the crura towards the other side. 
Others run towards the crus of the same side and apparently 
into it. 

(3) Oailar. Fibers seem to end in the lateral mass of the 
ocular lobe, numbers of them connect this with the more median 
spherical "punktsubstanz." Fibers connect these two masses 
and fibers from the surrounding cell groups run into one or 
both of them. 

Fibers connect the ventral epithelial-like cell region with 
the lateral dark mass, and also with an adjoining group of small 
cells more medially situated. 

252 Annals Entomological Society of America [Vol. IV, 

The more median ventral mass of epithelial-like cells lying 
ventral to and slightly out from the spherical fibrous area 
probably has connections with the larger more dorsal cells. 

The more dorsal of epithelial- like cells which are one side 
of a small cavity, send fibers into the lateral optic "punktsub- 
stanz" and are connected by fibers with the more central dorsal 
cells which adjoin it. 

(4) The Crura cerebri. The main mass of fibers goes up to 
the central portion of the "punktsubstanz" on each side and 
receives branches from all parts of it and also especially from 
the median and lateral dorsal cell groups and probably also 
from lateral masses. A large part of the band runs ventrally 
and forms or is joined into a commissure with fibers from the 
other side. Fibers come into this last from all ventral parts of 
the ganglion from both directions, from ventral parts of the 
mushroom bodies, and from various lobes of the deep staining 
mass of the ganglion. 

(5) Commissures. Two commissures have been mentioned, 
a ventral which was described above. The other more medial 
is found toward the caudal region. It is short and broad. 


1. In the abdominal ganglia practically all the nerve trunks 
and branches are mixed motor and sensory. 

In the thoracic ganglia, the three main trunks are mixed. 
Pure sensory divisions were found in connection with cephalic 
branches. The branches given off between the thoracic gan- 
glia and between the 1st thoracic and the subespohageal seem 
to be motor. At least the more caudal ones were determined 
to be, with fibers ascending and descending from the ganglia 
below and above. 

The three large nerve trunks connected with the subeso- 
phageal ganglion seem to' be mixed motor and sensory. The 
other smaller ones were not determined, but the small gustatory 
nerves were motor at least. 

The three main trunks connected with the supraesophageal 
ganglion were all mixed but the ocular. 

2. Sensory tracts were recognized entering thoracic and 
abdominal ganglia and distributed to various parts of the gan- 
glion to which they were connected. That is distributed to 
one or both sides either as individual fibers or as branches from 

1911] Nervous System of Corydalis 253 

one. Some tracts were found distributed to the next center 
above as well as the nerve center to which their nerve trunks 
were connected. Others were found passing in to the ganglion 
to which the trunk was attached but giving no branches and 
passing on up to end in the next above or higher up. 

A number of fibers apparently sensory were found passing 
through a number of ganglia without branches. These were 
often larger strands and although not traced as far as the head 
there was no reason to doubt that some of them were that long. 

Sensory fibers and tracts were however traced into the large 
head ganglia and were found distributed to all portions of the 
fibrous mass. Sensory tracts were easily traced in connection 
with the ocular trunk, connecting it with the special lateral 
masses of fibrous substance. 

.3. Motor tracts could be traced as having their origin from 
cells in the ganglion to which the motor trunks were connected. 
Motor fibers could be traced out of the ganglia some distance 
but were not followed in as much detail as the sensory. 

4. Association and descending tracts were recognized in 
all centers and at all levels of variable extent. Probably some 
of these were motor tracts. 

5. Cells of many sizes were found in all centers. The 
largest and the smallest functional nerve cells seemed to be for 
association, those of the former sort having extensive aboriza- 
tions, the second kind being much less extensively branched. 
Many of the medium sized cells were found to be motor. The 
general type was uni- or bipolar with one long branch which 
might run out for a considerable distance. Some association 
cells seemed to be bipolar with long processes running out in 
both directions. Some association cells seemed to be uni- or 
bipolar with all the branches coming off and branching again not 
far from the cell. A few multipolar forms were seen. 

Neuroblasts were found to some extent in all ganglia, but 
great masses of them were especially noticeable in the brain. 

Neuroglia networks filled in places under the chitin where 
there were no nerve cells and also formed more or less of a net- 
work in the regions where they were present. 

6. Cells were grouped about the central fibrous mass in all 
of the ganglia. In the abdominal they were found to be espec- 
ially abundant in the caudal and ventral regions, although the 
cell masses extended out laterally in all, and there were 

254 Annals Entomological Society of America [Vol. IV, 

often well marked mid-dorsal masses of large and smaller cells, 
as well as scattered cells in all parts. In the more cephalic 
ganglia, the cells are not quite the same in distribution. Cell 
groups and masses are found abundantly ventrally, but also on 
the dorsal and lateral sides and also great masses of them at the 
cephalic as well as the caudal end. 

In the supra esophageal ganglion, cells are especially abun- 
dant dorsally and on the cephalic and caudal borders, less abun- 
dant laterally and ventrally. The cells are very numerous. 
Each side of the middle line a group was located with very 
long processes, the cells being mostly large. 

Other groups were densely massed lateral to these, both 
cephalad and caudally, some of them were large, others small, 
and two groups of small cells on each side out near the optic 
nerves were found with a cavity near them. 

Nerve cells were found partly surrounding two masses of 
fibers On each side near the optic nerves. 

From the distribution of the cells and fibers there was no 
evidence of the dorsal cells being especially motor and of 
a ventral, particularly sensory region as Binet was led to 
think from experiments. I would rather incline to Kenyon's 
idea of ventral motor and dorsal sensory if I choose either of 
the two views, as undoubtedly most of the ventral cells in the 
thoracic and abdominal ganglia are motor cells, while many if 
not most of the fewer dorsal cells of these ganglia seem to be 
association cells, sending their fibers superficially over the sur- 
face or deeply into and through the fibrous mass to be associated 
with the cells and fibers of the ventral side. 

7. In all of the ganglia the central mass into which many 
of the nerve cells send their processes has a very complex ar- 
rangement of fibers and nerve terminations. Certain portions 
of this mass in all are much denser than the rest. 

In the brain this central "punktsubstanz" is somewhat 
more intricate in its texture on the dorsal side than on the 
ventral and is more or less lobed while off from the central mass 
there are two smaller groups of "punktsubstanz" in connection 
with the optic nerves on each side. In the central part of the 
brain is a stalk and root of a mushroom body on either side, but 
the cup is not present. The ventral portion of the ganglion is 
mostly made up of straight fibers. 

In all of the lower ganglia there are at almost any level, 
from one to three commissures recognizable, a dorsal, a median 

1911] Nervous System of Corydalis 255 

and a ventral. These are broken up at various levels so that 
there may be several, three or more, cephalic or caudal parts 
of these commissures. There are more in the 8th abdominal 
and in the subesophageal than in the others. 

From dorsal to ventral sides also, fibers chiefly from dorsal 
cells connect the upper and lower surfaces. 

In the brain there are two commissures a broad short median 
and a longer ventral. Many other cross and longitudinal 
fibers connect all levels. 

Cornell Univ., Ithaca, N. Y. 


Bauer, V. 1904. Zur innerin Metamorphose des Centralnervensystem der 

Insecten. Zool. Jahbr. T. XX. 
Benedicenti, A., 1895. Recherches histologiques sur le systeme nerveux central 

eperiphdriquc du Bombyx mod. Archiv. Ital. Biol. T 24. 
Berlese, A., 1907. Gli Insetti Vol Primo. 
Blnet, A., 1894. Contribution a I'^tude du systeme nerveux sousintestinale des 

Berthe, A., 1985. Studien uber das Centralnervensystem von Carcinus maenus 

Archiv. f. Milr. Anat. XLIV. 
Floyd, R., 1903. Contribution to the Nervous Cytology of Periplaneta orientalis. 

Mark Anniversary Volume. 
Faivre, E., 1857. Du cervaeu des Dytisquis consid^re dans ses rapports avec la 

locomotion. Ann. de Sc. Natur. Zool. ser. 4 T 8. 
Haller, B. 1904. Ueber den allgemeinen Bauplan des Tracheatensyncerebrums. 

Archiv. f. Mic. Anat. 
Hammar, A., 1908. On the Nervous System of the Larva of Corydalis cornuta L. 

Ann. Ent. Soc. of America. Vol. 1, No. 2. 
Hilton, W. A., 1902. The Body Sense Hairs of Lepidopterous Larvae. Am. Nat. 

Vol. XXXVI. 
Hilton, W. A., 1909. The Tracheal Supply in the Central Nervous System of the 

Larva of Corydalis Cornuta 1. Ann. Ent. Soc. of America. Vol. II, No. 1. 
Kenyon, F. G., 1896. The Meaning of the So-called "Mushroom Bodies" of the 

Hcxapod Brain. Am. Nat. Vol. XXX. 
Kenyon, F. G., 1896. The Brain of the Bee. Journ. Comp. Neurol. Vol. VI. 
Krauss, W. L., 1984. On the Nervous System of the Head of the Larva of Cory- 
dalis cornuta I. Psyche, Vol. IV. 
Newton, E. F., 1894. On the Brain of the Cockroach Blatta orientalis. Quart 

Tour. Mic. Sc. Ncu. Ser. Vol. XIX. 
Packard, A. S., 1880. The Brain of the Locust. Second Report of the U. S. 

Entom. Com. 
Saint-Remy, 1890. Contribution a I'^tude du cerveau chez les Arthropodes tra- 

cheates. Archiv. Zool. Exper et Gen. 2d Ser., V, 4th Memoir. 
Villanes, A., 1886. IV Memoire. Le Cerveau de la Guepe (Vespa crabo et vul- 
garis). Ball. Soc. Phil., Paris. 7. e Ser. T, 9. 
1887. Sur la morphlogie comparee du cerveau des Insectes et des Crustac^s 

(C. R. Acad. Scienc.) 
1887. Etudes histologiques et organologiques sur les centres nerveux et les 

organes des sens des animaux articul^s. Annales d. Sc. natur. Zoologie, 

7. e Ser. T. 4. 
1887. V Memoire. I. Le cerveau du Criquet (Oedipoda caerubscens et Calop- 

tenus italicus). Annales d. Sc. Natur. Zoologie 7. e Ser. T. 4. 
1893. VI Mem. II. Apercu general de 1' organization du systfeme nerveus des 

articul^s. Ann. Sc. Nat. (7) Vol. XIV. 

256 Annals Entomological Society of America [Vol. IV, 

A series of four photographs of cross sections through the 1st thoracic ganglion 
of Corydalis through a caudal level. The last figure is through the more central 
portion of the ganglion. Nerve cells show at the sides. The central fibrous mass 
is shown in all, and longitudinal fibers can be traced. In the last three figures 
cross commissures may be seen. The dark mass at the left or ventral side is part 
of the surrounding tissues and not a part of the ganglion. x60. 

Fig. I. Photograph of a cross section parallel with the long axis of the brain 
through its central region. One half only shown. The dorsal side is up. The 
optic nerve is the swelling off at the right above, while leading off below is the 
broad crus. The dorsal masses of cells show, also central fibers and the stalk and 
root of the mushroom body. x75. 

Fig. 2. A similar photograph from the same series cut farther one side, 
cephalad. x75. 

Fig. 3. Photographs of the larger cavity shown in Figs. 1 and 2. x250. 

Fig. 4. Section of the brain cut in a similar way as Figs. 1 and 2, but farther 
cephalad. The ventral side is at the left. The dorsal to the right. x75. 

Fig. 5. Longitudinal section of the sub-esophageal ganglion. Above the first 
branch to the left is one of the crura cerebri, the next it is the mandibular. Below 
the branch cut only through its edge is one of the ventral connectives. x75. 

Annals E. S. A. 

Vol. IV, Plaxe XV. 

ir. A. Uillun. 

Annai,'* E. S. a. 

Vol. IV, Plate XVI. 

ir. .1. nuiuii. 


William T. M. Forbes, New Brunswick, N. J. 

This paper may be considered a continuation of that pub- 
lished in the Annals of the Entomological Society of 
America, III, 94-132, with plates 10 to 20. The references to 
figures with the prefixed numeral I, refer to plates 13 and 14 of 
that article. 

In the caterpillar stage the Sphingidae may be defined as 
follows: With secondary hair on the body, epicrania, front, 
clypeus, maxillae and labium, but never on the antennae or 
distal parts of the maxillae and labium. There is almost 
always secondary hair on the adfrontals. It is present on the 
mandibles in Cressonia only, and the labrum bears a single 
additional hair in Pachysphinx. Front not more than half the 
height of the head (measuring here and elsewhere in this paper 
from the base of the front to the vertex. The frontal punctures 
are close together, and when distinct the primary setae are 
somewhat further from them than they are from each other, 
but they are distant from the outer edge of the front. The 
lobes at the two lower outer angles of the front tend to be large. 
Adfrontals not very wide, often narrow, not extending between 
the clypeus and mandible when distinct in the lower part; 
puncture a little below the upper seta. Clypeus narrow in the 
middle, often grading into the membrane below. Labrum well 
marked, often with very deep notch; vi distant from the margin, 
and usually directly below ii. Mandible with a large scrobe, 
bearing the two usual setae, one at the tip, and the other 
about half way out, toward the posterior side. Antennae with 
second joint most often twice as long as the first; the remaining 
joints exceptionally minute. Maxillary palpi stout, the second 
joint somewhat shorter than the first; submenta ill-defined, and 
mentum broad at the base. Spinneret broad, flat and short, 
with the usual three sclerites; labial palpi similar in form, and 
set obliquely to it, forming with it a sort of scoop (which would 
seem more useful in feeding than in spinning). 

Claws not distinctly notched, broadening abruptly at their 
base. Prolegs with a double row of hooks, about 20 to 60 in 
number; not very regularly arranged in Hemaris, etc. The 
hair on the prolegs is never as rudimentary as higher on the 


262 Annals Entomological Society oj America [Vol. IV, 

Secondary hair always minute, rudimentary, mostly visible 
only under a lens; but the body is often roughened by the 
tubercles; primary hair (iii and v alone are easy to identify but 
in Proserpinus flavoj'asciata i and ii are marked by larger black 
spots) often considerably better developed; but their tubercles 
are never raised, even when the others are. Segments with 
8 or 9 annulets, usually ill-defined in front. Usually with a 
wart, horn or peculiar marking on the dorsum of the eighth 
abdominal segment — if a horn, bearing tubercles and secondary 
hair. With other armature only in Ceratomia amyntor, Lint- 
neria eremitus and some exotic Smerinthinae. 

Tree-feeders with a few exceptions (L. eremitus, Deilephila, 
Protoparce, Choerocampa) or feeders on vines. 

In synoptic form the Sphingidae are distinguished as 
follows : 

1. Maxillary palpi three-jointed; the four upper ocelli in a rectangle; the upper 
setae of thoracic segments on a level JUGATAE 

1. Maxillary palpi with but two free joints; the four upper ocelli in a curve; the 

upper setEe of the thorax, when single, one above the other.. (FRENATAE) 

2. With outer hooks on the prolegs Microlepidoptera, etc., etc. 

2. With a single inner band of hooks on the prolegs 3 

3. Hooks of prolegs alternately of three lengths Butterflies 

3. Hooks of prolegs all of the same length 

Noctuidae, Notodontidae, Arctiidae, Eucleidae, etc. 

3. Hooks of prolegs alternately and regularly of two lengths 4 

4. Secondary hair present on antennse and palpi Lasiocampidae 

4. No secondary hair on antennae or palpi 5 

5. Secondary hair if present, confined to leg-bases .. Geometridae, Thyatiridae 
5. With dense secondary hair on body and head, including labrum 

Apatelodes (Eupterotidae ?) 

5. With considerable secondary hair on body and head, but little or none on 

labrum 6 

6. Primary tubercles represented by branching spines, or warts bearing several 

hairs Saturnoldea 

6. Primary tubercles bearing minute simple hairs, hardly distinguishable from the 
secondaries; and all hair minute SPHINGIDAE 

Semanophoile and Asemanophor.^ are separated rather 
by different tendencies than by sharp differentiating characters, 
and the same is true to an even greater extent of the subfamilies 
of each. Marking in a general way the Asemanophorce there 
may be mentioned the strong tendency for the head to taper 
toward the vertex (which none of the Semanophorce show) ; the 
densely granulated skin of the middle stages (except Lapara, 
and shared by one or two Semanophorce); the generally higher 
front, with more tendency to develop lobes at the lower outer 
angles. The first abdominal segment is never swollen, and the 
horn is never replaced by an eyespot; the clypeus tends to be 
wider, at least at the ends. 

1911] Study of Caterpillars of Sphingidae 263 

In the majority of the Semanophorce the last thoracic or 
first abdominal segment is much swollen, and the horn is fre- 
quently replaced by an eyespot. The front often makes a 
perfect equilateral triangle. The skin is very sparsely if at all 
granulated in the last stage, and is rarely granulated in the 
earlier stages. On the labrum, seta iii is quite generally on a 
level with ii, in the Semanophoras more often nearer the level 
■of vi, at least in normally formed labra. Normal oblique 
stripes are rather rare. 

If we except Pachysphinx occidentalis we might define the 
Smerinthinae as caterpillars with heavily granulated body in 
the last stage, and more or less granulated head, with the setae 
rising from the apex of the granulations. All except Pachy- 
sphinx have an acute triangular head, and even in this the head 
tapers strongly toward the vertex. The labrum is always nor- 
mal in arrangement of its setae, with the distance between the 
setae ii full half the width of the labrum, and with moderate, 
flaring notch. Except in its smooth skin, and normal horn 
Pachysphinx occidentalis agrees exactly with P. modesta, and in 
horn it agrees with Smerinthus. 

The Sphinginae, which comprise the rest of the Asemano- 
phorcB, are less homogeneous. Ceratomia has a densely gran- 
ulated skin, but is distinguished from all the Smerinthince by 
possessing a row of middorsal granulations. In the more 
specialized forms the labral setae are closer together around the 
notch, and this reaches its extreme in Cocytius, which has a head 
of normal Smerinthid form. Lapara, also with a triangular 
head, is easily separated from the Smerinthinae and placed in 
this series, as its labrum. (as well as markings and habits) 
agrees closely with Hyloicus. 

I cannot distinguish the three subfamilies of Semanophorce, 
■even by tendencies, and am inclined to treat them as a single 
subfamily. The eyespot in place of a horn occurs only in the 
Philatnpelince, but they are not all of the same type. Humped 
caterpillars occur in all three (Choerocampa, Darapsa, etc., 
Erynnyis) and cylindrical ones also (Deilephila, Sphecodina, 
Pseudosphinx) ; reduced horns (Pergesa, Pholus vitis, and 
Erynnyis) and normal ones (Xylophanes, Darapsa, Hemaris) ; 
rounded heads (Deilephila, Macroglossa, Pseudosphinx) and 
squarish ones (Choerocampa, Darapsa, Erynnyis). The labrum 
•of Pseudosphinx and Erynnyis is a little peculiar, but that of 

264 Annals Entomological Society of America [Vol. IV, 

Deilephila comes as near as that of Hemaris, to it. Hemaris 
croatica is an almost perfect connecting link between Hemaris 
and Macroglossa. Altogether a tabulation of the genera which 
shall be workable must be mainly artificial in its arrangement. 


1. Head high and triangular, no horn Lapara 

1. Head rounded, or horn more or less developed 2 

2. Head half higher than wide, horn well defined, enlarged tubercles on anal 

plate 3 

2. Otherwise 4 

3. Four tubercles on anal plate, face smooth (Dilina)* 

3. Two tubercles on anal plate, face rough Cressonia 

4. Head much higher than wide and triangular; horn rather small and soft, 

not well distinguished from the body 5 

4. Head about as wide as high, or trapezoidal 6- 

5. Face smooth, sides tuberculate (Amorpha) 

5. Face as rough as sides of head, all heavily tuberculate 

Smerinthus and Paonias 

6. Seven setae on labrum, transverse ridges, — one each on meso- and meta- 

thorax Pachysphinx 

6. Six setae on labrum, transverse ridges more numerous or wanting 7 

7. Body heavily granular, with granular obliques, middorsal line, and subdorsal 

on thorax, the latter raised into two pair of short horns Ceratomia 

7. Body smooth or nearly so in last stage, no middorsal granules 8 

8. Labrum with ii decidedly higher than i Hyloicus (cupressi?) 

8. Labrum with ii on a level with i or lower 9 

, 9. Head triangular, smooth, as well as cervical shield; anal plates exceptionally 

rough; labrum with a deep narrow notch, with the setae i, ii and vi crowded 
around it Cocytius 

9. Head rounded on the vertex, but in Chlaenogramma with two enlarged granu- 

lations; anal plate rarely rough; setae ii of labrum almost half as far apart 

as width of labrum , or more 10 

10. Head much higher than wide, and tapering to about half its width at the 
vertex 11 

10. Head as wide as high; tapering decidedly toward the vertex 12 

,10. Head nearly as wide as high, rounded or squarish, or if tapering a little then 

strongly granulated 17 

11. Horn nearly twice as long as height of head Chlaenograinma 

11. Horn slender, and but little longer than height of head. Daremma (undulosa) 

12. Horn recurved at tip (Acherontla) 

12. Horn regularly pointed 13 

13. Notch very shallow, in depth only 1-7 height of labrum .... Phlegethontius 

13. Notch of labrum at least twice as deep 14 

14. Horn very slender, head well rounded at the sides Daremma (catalpa) 

14. Horn normal or short 15 

15. Mesothorax acutely humped, sometimes surmounted with a tubercle .... 


15. Body normal in form 16 

16. Head normally with two pair of back side-stripes Herse 

16. Head normally with one pair of dark side-stripes Sphinx 

17. Head heavily granulated, body cylindrical, with normal obliques on sides. . IS 

17. Head not granulated, or body swollen, or without oblique stripes but with a 

continuous subdorsal picked out in enlarged granulations 19 

18. Body somewhat granulated in last stage, especially on the obliques, etc. 

18. Body unusually smooth Dolba 

* Exotic genera in parenthesis. 

1911] Study of Caterpillars of Sp>hingidae 265 

19. Setae i, ii and iii of the labrum on a level; labral notch very shallow, abdo- 
men cylindrical and rather slender 20 

19. Setae i higher than ii and iii, ii usually decidedly higher than iii 21 

20. Metathorax swollen, supraanal not noticeably armed Erynnyis 

20. Cylindrical, supraanal with two large tubercles Pseudosphinx 

21. Horn reduced to a granule or replaced by an eyespot; metathorax strongly 

swollen; head full as wide as high and squarish Pholus 

21. Horn present in all forms with swollen body, swelling rather belonging to the 

first segment of the abdomen than to the thorax; head rounded, or if 
squarish, full as high as wide 22 

22. Horn wanting (Deilephila vespertilio) 

22. Horn replaced by an eyespot 23 

22. Horn conical well developed 24 

23. Head very rough in last stage, horn very slender in middle stages, replaced 

by a high tubercle in the next to last Sphecodlna 

23. Head nearly smooth in the last stage; horn stout in penultimate 

Proserpinus (in part) 

24. Horn blunt and cylindrical; body marked with a large subdorsal eyespot on 

the thorax, and with well defined distant annulations, beginning with 
the second abdominal segment (Daphnis) 

24. Horn acute; no eyespot on side of thorax 25 

26. Body more or less swollen on first segment of abdomen, thence tapering to 

the head 30 

25. Body not swollen 26 

26. Head and general surface of body distinctly but sparsely tuberculate, the 

setae rising from the apex of the tubercles; cervical and anal shields 
rougher than the rest of the body 27 

26. Body smooth, head rarely with very slight tubercles, from the slope of which 

the setas rise 28 

27. Tubercles only visible under a lens (Macroglossa) 

27. Head and cervical shield appearing rough to the naked eye Hemarls 

28. Notch of labrum hardly 1-3 its height; setEe i and ii spaced about in the ratio 

2:3; vi only 1-3 way up from the apex of the lobe toward ii and much nearer 
the middle line Deilephila 

28. Notch of labrum deeper, setae i and ii only about half as far apart as the setae 

ii are from each other; vi nearer to ii than to the apex of the lobes. . . .29 

29. Front about half as high as height of epicrania; without normal oblique stripes 

(with reversed oblique spots) Proserpinus (in part) 

29. Front smaller; with seven normal obliques 14 

30. Horn not so long as height of head 31 

30. Horn very slender . '. (Hippotion) 

30. Horn normal 32 

31. Head very finely granular, appearing under low power smooth and dull 


31. Head distinctly vermiculate Amphion 

32. Head sparsely tuberculate 33 

32. Head smooth and dull under low power, granular under high power, with 

smooth areas about the setae 34 

33. Body slightly swollen, spiracles pale with two black bars Deidamia 

33. Body much swollen, spiracles dark with a white dot at each end 

Darapsa, Ampeloeca, (Clarina) 

34. Front higher than wide (Theretra) 

34. Front wider than high Xylophanes 

Sphinx. {Hyloicus in part). Head slightly tapering toward 
the top, or with sides rounded out {S. drupiferarum), with 
irregular vermiculate grooving; the setae rising as often from 
the grooves as from the elevated portions. Front about \ 
height of head, with lobes at the lower outer angles well marked 

266 Annals Entomological Society of America [Vol. IV, 

and about \ to \ its height. Ends of clypeus as wide as 
^ height of front. Labrum high, with a notch \ to \ its 
width, the setae i and ii spaced about in the ratio 1 :2, vi a 
little nearer in 5. drupiferarum, a little farther off in 5. gordius, 
luscitiosa and eremitus; iii, a little higher than vi but much 
below ii; notch deep and slit-like in S. gordius and luscitiosa, 
shallower and more flaring in the others. Mandible with a few 
(perhaps four, but ill defined) large teeth. Second joint of 
antenna decidedly longer than the first, and nearly twice as 
long as wide. First segment of body much larger in diameter 
than head and more or less enclosing the back of it. Skin 
entirely smooth, granulated until the last stage; supraanal 
more or less granulated. Horn normal, much longer than head, 
and curved downward. The seven stripes similar. 

There is also some variation in the horn. In the majority 
of species it is cylindrical in the basal part, and strongly down- 
curved in its entire length; but in chersis the basal part is more 
often nearly straight, and in gordius it is regularly conical and 
the whole horn is almost straight. The European species 
ligustri, which comes next to drupiferarum in the adult, in the 
caterpillar resembles it closely in markings, but has the normal 
Sphinx head. (Fig. 1). That of drupiferarum would not differ 
in face view from Fig. 10. See also I, Figs. 39-41 of S. gordius. 

Lintneria has a conical hump on the mfesothorax, which in 
the next to last stage, and sometimes in the last, is surmounted 
with a hard tubercle. Otherwise it resembles Sphinx in struc- 
ture. It is considered a subgenus of Sphinx. (L. eremitus, 
Fig. 8.) 

Hyloicus. Head rounded and decidledy larger in diameter 
than the body. Horn straight and slender. Labrum with 
very shallow widely flaring notch; with setae i and ii nearly 
evenly spaced; ii much lower than i and the setae i, ii and vi 
of each side forming an equilateral triangle. Second joint of 
antennae only half longer than wide, and first joint very short. 
Supraanal long and triangular. Fig. 10. 

//. pinastri is longitudinally striped with green and white, 
with a broad red dorsal, or else suffused with red. Horn black. 
H. cupressi of the southern states is reported as similar, with 
the white lateral stripes broken into patches; and is probably 
similar in structure. 

1911] Study of Caterpillars of Sphingidae 267 

Herse. I cannot distinguish Herse from Sphinx by any 
satisfactory characters. The head in both convolvuli and 
cingulata is intermediate between those of drupijerartim and 
the other Sphinxes. Setae i and ii of the labrum are about f 
as far apart as the distance between the two setae i, and i is 
but Httle higher than ii. The two main joints of the antennae 
are practically equal. In the only specimen of H. cingulata I 
have seen the horn is very short, but this may be an abnormal- 
ity; it is normal in H. convolvuli. 

Dolba. The head does not taper decidedly toward the top, 
and is decidedly granular, the setag rising from the apex of the 
widely separated granules, as in Smerinthus. Otherwise it 
agrees with those species of Sphinx in which the labrum is not 
deeply notched. The head comes surprisingly close to that of 
Darapsa, but may be distinguished by the decidedly higher front, 
and the fact that iii of the labrum, as in most Asemanophorae, 
is nearer to the level of vi than ii. I, Figs. 39-41 represent not 
this species but Sphinx gordius. 

Atreides has a very similar head. The supraanal plate is an 
equilateral triangle; the markings are picked out with raised 
granules, and there are several transverse rows on the thorax, 
and scattered granules on the abdomen as in Smerinthus, but 
very widely scattered and small. 

Acherontia. Has a slight transverse hump on the meso- 
thorax (suggested in some Sphinx) ; the tip of the horn is recurved 
sharply; the fine annulations are wanting from the thorax. 
Otherwise entirely like Sphinx (e. g. S. kalmiae). A. atropos 
examined, European. 

Ceratomia. (sens, str.) Head about as wide as high, decidedly 
tapering, somewhat granulated in back, but with the setae not 
springing from the granulations, structurally as in the lower 
species of Sphinx. Body densely granulated, with the sub- 
dorsal and obliques picked out in raised granulations, and also 
with a mid-dorsal row. The subdorsal row is produced on the 
meso-, and metathorax into two pairs of short soft horns. 
(I, Figs. 45 and 49.) Aside from subfamily characters, I have 
seen no trace of kinship between this species and Daremma. 
The latter seems to come closer to Chlaenogramma. 

Daremma. Skin smooth; horn normal, rather slender and 
short; or longer but very slender (catalpce). Head slightly 
granular, but the setae do not rise from the apex of the gran- 

268 Annals Entomological Society of America [Vol. IV, 

ulations. Otherwise the characters common to undulosa and 
catalpcB are shared by Sphinx. 

In D. undulosa, the type, the head is \ higher than wide, 
very strongly tapering toward the vertex, the horn is only a 
little slenderer than normal, and the labrum is moderately 
notched, with seta iii in the normal place. In D. catalpce 
(Fig. 11), the head is broad and rounded, shaped as in Deil- 
ephila; the labrum is quite deeply notched, with iii nearly on a 
level with ii, as in the Semanophorae; the body is unusualty 
cylindrical and the horn is very slender. I have seen a spec- 
imen from the U. S. National Museum, with no data but the 
name Daremma hageni. It is a typical Smerinthus in structure, 
and could be the fourth stage of one of the larger species 
(cerisyi), but is rather sparsely granulated. I should suspect 
it was misidentified. 

Chlcenogramma. Head like that of Daremma undulosa, but 
with inconspicuous enlarged granules on the vertices. Body 
like Atreides plebius. Horn normal in length. 

Cocytiiis (antceus). Head not at all tuberculate, but decid- 
edly higher than wide, and terminating in two enlarged gran- 
ules, like a fourth-stage Smerinthus, but higher and closer 
together at the vertex. Front full as high as wide, the lobes at 
the outer lower angles are not only well marked in outline, but 
project very conspicuously; front less than | height of epi- 
crania, iii and ii of the labrum are on a level, but the notch runs 
even higher, and is very narrow. The distance between the 
two setae ii is less than § the width of the labrum, and i, ii 
and vi are all located practically in the notch. Cervical shield 
as smooth as head, the anal is very rough, like the horn, and is 
a narrow triangle. Skin not at all granulated. Altogether an 
unusually distinct genus for this series. The structure so far 
as it is not unique suggests Sphinx rather than any other 
genus, but I understand the moth comes nearer to Phlege- 

Phlegethontius {Protoparce) Notch of labrum only one- 
seventh its height, not reaching the level of seta vi; iii and iv 
not so high as usual. Caterpillar distinctive in appearance, 
but not otherwise separated in structure from normal Sphinx. 
S. rusticus is said to be sparsely granulated on the lines. I, 
Fig. 51. 

1911] Study of Caterpillars of Sphingidae 269 

Lapara (Ellema). Skin quite smooth, not only in the last, 
but in earlier stages. Horn wanting entirely. Anal plate 
fully as long as wide and acute. Head somewhat higher than 
wide, (Fig. 4), in the earlier stages extremely high, triangular, 
with small and distant tubercles, like Fig. 12; front | 
its height, higher than wide. Labrum (Fig. 5) with a very 
shallow notch, broadly flaring, with the apex of the lobes far 
to one side and the outer edges nearly straight; i lower than 
ii, but not so much so as in Hyloidus, i and ii about equally 
spaced; both crowded down toward the margin; vi decidedly 
nearer the middle line than ii. Scrobe of mandibles smaller 
than usual. Second joint of antenna hardly longer than wide, 
and first joint very short. First ocellus directly behind the 
second, and nearer to the posterior one than the second is to 
the fourth. With longitudinal stripes, or checkered, never 
with obliques. 

Except for the labrum and markings, which are essentially 
as in liyloicus pinastri, there is nothing to connect this genus 
with the Sphinginae in the caterpillar; there are a couple ot 
parallelisms to the Smerintliince; the shape of the head, and low 
first ocellus as in Cressonia. 

I cannot distinguish the species in the caterpillar. 

Smerinthus (Sphinx) and Paonias {Calasymboliis) (I, Figs. 
42-44). Head decidedly higher than wide, triangular; with 
nearly acute apex and sides somewhat rounded out; with 
numerous widely spaced raised tubercles, each bearing a seta; 
front about as in Sphinx, with several tubercles somewhat 
smaller than those on the epicrania; labrum with a notch about 
34 its . width, in depth, with the setae arranged as in 
Sphinx, but the distance between the two setae vi is full half 
the width of the labrum (in the Sphinginae it is mostly dis- 
tinctly less than half) ; iii, iv and v about equally spaced on the 
outer edge. Clypeus and mandible and antennae as in Sphinx, 
but the adfrontals are somewhat wider. Body finely gran- 
ulated, strongly tapering toward the head; first prolegs less 
used than the others and sHghtly reduced. Subdorsals on 
thorax, and obliques on abdomen marked by rows of raised 
granules. No granules on the middorsal line, but they show a 
tendency to arrange themselves in a row on each side of it. 
Horn soft, not well distinguished from the body and about as 
long as the height of the head, not down-curved. Supraanal an 

270 Annals Entomological Society of America [Vol, IV, 

equilateral triangle in shape, not specially armed, acute; 
The transverse rows of granulations on the thorax are all about 
equal. Ocelli in normal arrangement, as in Sphinx. 

The species show very little distinctive in the way of struc- 
ture (or for that matter in color and markings) P. (Calasym- 
boliis) astylus may be a' little rougher, with better developed 
setae than the others, both on head and body, and the horn 
seems a little better defined in S. ocellatus than the others. 

Pachysphinx {Triptogott, Marumba). Head wider than high, 
with sparse granulation on the sides, nearly smooth, but a 
little vermiculate on the face; the apex bluntly rounded. No 
subdorsal row of granulations on the thorax but the meso- and 
meta-thorax each have one high transverse crest. Horn soft, 
variable in size. The three lower ocelli form a right triangle, 
the posterior being unusually high. The labrum has an addi- 
tional seta, on the margin; the four marginal setag that result 
are about equally spaced. 

1. P. modesta. Body normally granulated, about as in 
Smerinthus; horn minute, about rs in. long; thoracic crests 
high and granular. (Fig. 7.) 

2. P. occidentalis. Body smooth, with a few raised gran- 
ules on the last oblique line, only; horn about as long as height 
of head. Thoracic crests rounded over and hardly distinguish- 
able. (In the penultimate stage it is granular like P. modesta.) 
This, as may be seen, is very different from the eastern form 
modesta. (Bred from the egg by Mr. Brehme; Western.) 

Amorpha {A. populi of Europe) Labrum very deeply 
notched (like Cressonia). Head decidedly higher than wide, 
triangular, smooth on the face. Anal plate unarmed, and 
horn, etc., as in Smerinthus. The last four oblique rows of 
granules very distinctly extend over three segments. The 
characters are nicely intermediate between Smerinthus and 
Dilina, showing no special closeness to Pachysphinx. 

Dilina {Mimas). Horn sharply separated from the body and 
down curved as in Sphinx; half longer than width of head, 
mostly cylindrical. With a longitudinal subdorsal row of 
granules on the thorax. Head half higher than wide, and 
acute-triangular. Face smooth, the sides of the head sparsely 
tuberculate. Supranaal with four raised tubercles in a rec- 
tangle. Otherwise about as in Smerinthus. D. tiliae of Europe. 

1911] Study of Caterpillars of Sphingidae 271 

Cressonia. With onlj'' one large pair of enlarged tubercles 
on the supraanal plate. Front as wide as high and only one- 
fourth as high as the epicrania. Entire head tuberculate; first 
ocellus moved down and posterior one up so that they are not so 
far apart and as the second and fourth are from each other. 
Mandibles ivith a tuft of secondary hair on the outer part of the 
scrobe. Otherwise as in Dilina. C. juglandis. (Figs. 12 
and 13.) 

Deilephila (Celerio) (Fig. 9, and I, Figs. 46 and 52). Head 
distinctly wider than high, rounded on top, roughened with 
irregular grooves, but not as strikingly as is usual in the 
Sphingince. Front triangular, the outer edge but little 
sinuous; clypeus narrower at the two ends than in the Sphin- 
gina. Labrum with a broadly flaring notch only one-third as 
deep as the height of the labrum or less, setae i and ii about two- 
thirds as far apart as the setae i are from each other, nearly on 
a level; vi less than half way up to ii. Antenna with the first 
and second joints nearly equal in length and diameter. Ocelli 
with the first four of nearly equal size, in a regular curve, but 
the second decidedly nearer to the third than to the first. 
Posterior ocellus about half way between the upper and the 
lower; the three lower ocelli forming approximately a right 
triangle. Skin smooth, the tubercles marked by tiny chitinous 
rings, those of the primaries two or three times as large as the 
others. Horn normal, moderate, entirely wanting in D. 
vespertilio. Supraanal broad. Caterpillar as a whole cylin- 
drical, tapering toward the head, almost always with a conspicu- 
ous row of subdorsal spots or eyespots, which are all about equal 
in size except the last. The front is smallest in D. euphorbice. 

D. litieata, gallii, euphorbiae, lathyri and vespertilio were 
examined. Aside from those mentioned above the differences 
come down to a slight variation in the depth of the notch of the 
labrum; and the markings. 

Chcerocampa (in the broader sense). (Fig. 6.) Head squar- 
ish, full as high as wide, very finely granular, so as to appear 
smooth and dull with low power; the region about the setae may 
be glossy but not raised. Front triangular, wider than high 
(except in alecto) at least |- height of head; posterior ocellus 
lower making the triangle formed by the three lower ocelli 
acute-angled; second joint of antenna about twice as long as 
first; notch of labrum at least 5 its height, usually more; i 

272 Annals Entomological Society of America [Vol. IV, 

and ii only half as far apart as the two setse i are from each 
other; vi nearer to ii than to the tip of the lobes. Body with the 
first abdominal segment much swollen and bearing an eyespot, 
with or without less prominent ones on the following segments; 
thorax lacking the mottling characteristic of the abdomen in the 
darker forms; horn regularly conical; skin smooth. 

Subgenus Theretra. Horn well developed; large eyespot 
normal, followed by a series of simpler ones in a subdorsal 
stripe; pattern of abdomen wanting from the first segment, as 
in some Philampelince; body with dark obliques slanting up 
and back. T. alecto. 

Subgenus Xylophanes. Horn and eyespots as in Theretra. 
Front distinctly wider than high; labral setae i and ii a trace 
farther apart and nearly on a level; notch broad and only \ 
height of labrum; second joint of antenna only half longer than 
the first. Perhaps a little near to Deilephila than the others 
are. X. tersa. 

Subgenus Pergesa. Horn very short and sharply down- 
curved; the first eyespot only is present, and it has a black 
shade below. Setse i of labrum much higher than ii, and thev 
are spaced as in Theretra, that is, in the ratio 1:2. Notch 
nearly half height of labrum; antennae with second segment 
twice as long as first. 

Subgenus Hippotioii. Similar to Pergesa, but with the horn 
long and slender (fig. 15). 

Pholiis (Philampelus). Horn reduced to the merest rudi- 
ment in vitis, replaced by an ej'espot which bears a pale cres- 
cent in its posterior part, in pandoras, achemou and labrusccc; 
said to be completely wanting in a Texan species. Metathorax 
much swollen, first segment of abdomen distinctly smaller. 
The front of the body is completely retractile in P. achemon 
and pandorus, apparently less so in vitis and labrnscae. Supra- 
anal broad, not specially armed. Head fully as wide as high, 
squarish; front higher than wide, and nearly half its height. 
Setae i, ii and vi of labrum especially close together. 

Ampeloeca. ' (Darapsa, Everyx, Ampelophaga). Head 
rounded, with widely separated small tubercles, similar to 
fig. 16, but higher; front more than ;= its height, decidedlj' 
higher than wide, also tuberculate. Labrum with a fairly deep 
notch; setae as in Choerocampa. Body decidedly swollen at first 
segment of abdomen, but not enough to withdraw the head. 

1911] Study of Caterpillars of Sphingidae 273 

Horn decidedly longer than head. With a subdorsal line, 
and the normal obliques. Spiracles red with white ends 
(agreeing with the Macroglossa group, Clarina, and some speci- 
mens of Protoparce). Otherwise like Choerocampa. 

In A. versicolor the horn is nearly twice as long as the head, 
thick and strongly down-curved; in myron it is shorter, straight 
and conical, but apparently it is exceptionally variable in all 
three species. 

Clarina. Horn only about as long as head; tubercles of 
head very small and inconspicuous; otherwise wholly like 
Ampeloeca, — with continuous subdorsal. C. syrtaca, of Syria. 

Deidamia. Horn rather longer than head. Supraanal an 
acute equilateral triangle. Spiracles pale with a black bar on 
each side. Front full as wide as high. Body but little swollen. 
Otherwise like Ampleoeca. 

C. syriaca and D. inscriptum are marked almost exactly 
like A. myron, but with evanescent obliques. 

Atnphion. Head irregularly roughened, without raised 
isolated tubercles, squarish, as in related genera. Supraanal 
as long as wide and acute; spiracles dark. Horn shorter than 
height of head. Front full as high as wide, third ocellus much 
enlarged (as in Clarina and Deidamia also). Body but little 
swollen. Otherwise like the related genera (I, figs. 48 and 53.) 

Sphecodina. Body not at all swollen on the first abdomi- 
nal segments. Head large, very rough in the last stage, but 
without raised tubercles; in the next to last as in Ampelceca. 
Front higher than wide, -i height of head. Clypeus broader 
than in Deilephila, etc. Labrum like Pergesa. Horn replaced 
by a wart ; in the last stage similar to that of Pholiis, in the next 
to last high, and cylindrical; before that surmounted by a slen- 
der horn, which rises, not as in Pholus from its posterior side, 
but from the middle. Supraanal wide. 

Proserpinus (Fig.- 3). Horn normal but rather short {gaurce) 
or replaced in the last stage by an eyespot, which may be nearly 
fiat (proserpina), or with an obliquely conical center (flavofas- 
ciata, juanita). Head and skin smooth. Spiracles single- 
colored, yellow in flavofasciata and proserpina, black in gaurce. 
Head squarish, higher than wide in proserpina, full as wide as 
high in flavofasciata. Adfrontals only about | as wide as 
front is high, with their setae i below the top of the front. Front 
nearly half as high as head, broadly triangular. Ocelli normal. 

274 Annals Entomological Society of America [Vol. IV, 

Labrum with a moderate or rather shallow notch, with the setae 
i and ii only half as far apart as the setae i are from each other 
(thus agreeing with the preceding genera rather than with 
Hemaris). Supraanal an equilateral triangle; joints of antennae 
nearly equal in diameter. In the next to last s,tage flavofasciata 
at least, has a horn similar to that in adult gaiirce. 

Macroglossa. Head and body nearly smooth, but marked 
by slight raised white tubercles. Head very small and round, 
the body sharply tapering toward it. Horn normal, longer 
than height of head. Adfrontals i height of front in width, 
and with seta i higher than the top of the front. Third, fourth 
and lower ocelli evenly spaced, and very close together, nearer to 
each other than to the posterior. Front t height of head ; dis- 
tance between setae i and ii of labrum | that between the 
two setae i. True legs single-colored; spiracles red with white 
spots at the two ends. The genus shows likenesses to Hemaris on 
the one hand especially to H. croatica, which is very similar; and 
to the Darapsa group on the other, rather than to Proserpinus. 

Hemaris (Haemorrhagia) . Head rough and tuberculate; 
cervical shield, etc., also rough, and skin generally with more or 
less distinct raised tubercles. Horn moderate, or long and 
slender. Front I height of head. Otherwise as in Macroglossa, 
even as to the coloring of the spiracles. The species I have seen 
make a very good graded series, from Macroglossa to //. thysbe. 

H. croatica. Cervical shield no rougher than head, without 
any distinct anterior ridge; lower ocellus close to the next one 
as in M. stellatariim. Legs without any black. Head regu- 
larly rounded, not distinctly higher than wide. 

H. rubens. Head and cervical shield with fine granulations 
separated from each other by about twice the diameter of a 
gramilation, except toward the anterior edge of the cervical 
shield, where they are nearly in contact, and in a single even row. 
Front, and head as a whole higher than croatica, agreeing with 
the following species. Feet with a little black on the front of 
the coxjB only. Horn rather short. 

H. diffinis (typical). Cervical shield with the granules 
no larger than in H. rubens, but with the surface of the shield 
raised into ridges, so as to appear much rougher, considerably 
rougher than the head. The granules on the anterior edge 
make a ridge, but they are not confluent and the ridge is not 
well defined. The true legs have the femora marked with 
deep black brown. 

1911] Study of Caterpillars of Sphiiigidae 275 

//. diffinis axillaris. Head conspicuously rough. Anterior 
band of cervical shield of crowded granulations, not all in a 
single row; the shield decidedly rougher than in the normal 
form; horn longer than head. 

H. thysbe. Even rougher than the preceding, the cervical 
shield with the granulations almost in contact, and on the 
anterior edge more or less confluent. Horn long and the outer 
part slender. Femora jet-black and very conspicuous. 

In the next to last stage, H. diffinis, at least can hardly be 
separated from Macroglossa. The horn is minute. 

Eriunyis (Dilophonola). Seta ii of labrum fully as high as i, 
and the setae i, ii and iii almost evenly spaced; iii, iv and v 
about evenly spaced along the outer edge. Supraanal sometimes 
with the rudiments of a pair of tubercles. Notch of labrum 
hardly | its height, in depth. Head squarish, and full as 
high as wide; metathorax sharply humped; the abdomen ab- 
ruptly smaller, and cylindrical. Horn short. In E. edwardsii 
the horn is somewhat shorter than the head, in E. ello, only a 
third as long, and only twice as long as thick. The tubercles 
on the anal plate are distincter in edwardsii, and the supraanal 
is narrower. 

Pseudosphinx (I, Figs. 47 and 50). Cylindrical with slender 
horn. Fourth ocellus as far from the lower as from the first; 
front wider than high and nearly half as high as head. Labrum 
very shallowly notched, with the setas i and ii nearly on a level, 
and almost evenly spaced; iv directly below iii and much nearer 
to it than to v. vi not far from margin. Two conical spines 
on anal plate. 

The following species were examined. I am especially 
indebted to Prof. J. B. Smith; Dr. Geo. Dimmock, Mr. William 
Beutenmuller, and Dr. H. G. Dyar, for the privilege of examin- 
ing specimens in their own collections and in those of the insti- 
tutions they represent. Exotic species are in italics; those from 
the far west, or from Florida only, are also indicated. 


Acherontia atropos 
Herse convolvuli 
Cocytius antceus Fla. 
Phlegethontius quinquemaculatus 

Carolina (sexta), I, Fig. .51. 
Atreides plebeius 
Dolba hylaeus 

276 Annals Entomological Society of America [Vol. IV, 

Spliinx ligustri II, Fig. 1. 




gordius, I, Figs. 39-41 

Lintneria eremitus, II, Fig. 8. 
Hyloicus pinastri, II, Fig. 10. 
Lapara bombycoides, II, Figs. 4-5. 

Chlcenogramma jasminearum 
Daremma undulosa 

catalpae, II, Fig. 11. 
hageni ? (W) 
Ceratomia amyntor, I,, Figs. 45 and 49. 


Pachysphinx modesta, II, Fig. 7 

occidentalis (W) 
Smerinthus ocellalus 

Paonias excsecatus 

myops, I, Figs. 42-44. 
Amorpha populi 
Dilina liliie 
Cressonia juglandis, II, Figs. 12 and 13. 


Deilephila lineata, II, Fig. 9. 

gallii, I, Figs. 46 and 52 



Ckceroratnpa {Pergesa) elpenor 

Charocampa (Hippotion) ceterio, II, Fig. 15. 
Chwrocampa (Therelra) alecto 
Choerocampa (Xylophanes) tersa 


Pholus achemon 


vitis (fasciatus) 

labruscae (Figure only) Fla. 

Daphnis nerii 
Clarina kotschyi syriuca 

Ampelophaga (Ampeloeca) myron versicolor 
Ampelophaga (Darapsa) choerilis 
Deidamia inscriptum 
Sphecodina abbotii 

Amphion nessus I, Figs. 48 and 53; II, Fig. 14. 
Proserpinus proserpina, II, Fig. 3 
Macroglossa stellatarnm 


Hemaris croatica II, Fig. 16 

rubens (W) 

diffinis and form axillaris 

Erynnyis alope edwardsii Fla. 

ello Fla. 
Pseudosphinx tetrio Fla., I, Figs. 47 and 50. 

lyll] Study of Caterpillars of Sphingidae 277 


A. Horn completely wanting, head half higher than wide and triangular 

Lapara coniferarum and bombycoides 
A. Eighth abdominal segment bears a little hard nodule (caterpillar brilliantly 

marked with black, white and red) Pholus fasciatus (vitis) 

A. Eighth abdominal segment with an eyespot, otherwise unarmed B 

A. Eighth abdominal segment with a horn, otherwise unarmed F 

A. Eighth abdominal segment with a horn, mesothorax sharply humped or tuber- 

culate Lintneria eremitus 

A. Eighth abdominal segment with a long horn, and thorax with four much short- 

er ones Ceratomia quadricomis 

B. Slanting white patches on the sides C 

B. Pale on the sides, obliquely mottled, eyespot white. .Pholus labruscae (Fla.) 

B. Otherwise marked; eyespot dark-ringed D 

C. White patches are full three times as long as broad Pholus achemon 

C White patches are 21.2 times as long as broad Pholus pandorus 

D. Checkered contrastingly with red and green Proserpinus juanita 

D. No red , E 

E. Eyespot ringed with jet-black Proserpinus flavofasciata 

E. Eyespot ringed with brown Sphecodlna abbotii 

F. Slanting lines on the sides, sloping upward toward the rear G 

F. Slanting lines, if present, running in the opposite direction Y 

G. Slanting lines dark and more than seven, with a double set on the dorsum 

Amphion nessus 

G. A single slanting line running up to the horn Cocytius antaeus 

G. Numerous slanting lines, on the dorsum only Cocytius cluentius 

G. Slanting lines indefinite in number or evanescent; spiracle marked with two 

vertical bars, caterpillar a little humped Deidamia inscriptum 

G. Slanting lines six or seven, and distant H 

H. Head rounded, with two pair of vertical dark stripes; on Convolvulaceae 

Herse cingulata 

H. Head mostly with one or no vertical dark stripes; on other plants I 

1 . Horn rudimentary Pachysphinx modesta 

I. Horn as long as head, or longer J 

J. Skin rough and granular, head triangular K 

j. Skin nearly smooth, but with the stripes marked with raised granulations. .0 

J. Skin smooth P 

K. Horn well defined and much longer than height of head. .Cressonia juglandis 
K. Horn about as long as height of head, and not very distinct from body, 

straight L 

L. The oblique stripes irregularly shaded with red patches, . , Paonias astylus 
L. One of two pairs of red spots alone, or with one or two much larger than the 

others Paonias myops 

L. With a number of equal red spots or with none M 

M. With none, horn normally blunt, or blueberry P. astylus 

M . Horn always acute N 

N. Horn pink, violet or blue Smerlnthus cerisyi and jamaicensis 

N. Horn normally blue-green Paonias excaecatus 

N. Horn normally yellow-green, yellow on the sides Paonias myops 

O. Head broad and rounded, granulated Atreides plebeius 

O. Head high and tapering, nearly smooth Chlaenogramma jasminearum 

O. Head broad and tapering, nearly smooth 

Phlegethontius rusticus, and occasional specimens of Sphinx chersis, etc. 
P. First segment of abdomen much swollen, subdorsal stripe well developed. .Q 
P. Caterpillar nearly cylindrical, subdorsal stripe present on thorax only....S 
Q. Subdorsal stripe complete Ampelophaga myron 

* See "Field Tables of Lepidoptera (1906) p. 69. 

278 Annals Entomological Society of America [Vol. IV, 

Q. Subdorsal stripe broken in the middle R 

R. Horn near twice as long as head, and down-curved A. versicolor 

R. Horn little longer than head and nearly straight A. choerilis 

S. With pale substigmatal bands on the segments which have obliques, meeting 

the obliques to form chevrons Phlegethontlus 5-maculatus 

S. No stigmatal band T 

T. With the obliques showing no trace of red and edged above with a row of 

black spots, horn reddish Phlegethontlus Carolina 

T. Obliques not edged above with a row of black dots; usuallv with red or violet. 

U. Horn little longer than the height of the head, which is decidedly higher than 

wide Daremma undulosa 

U. Horn about half longer than the head, which is as wide as high V 

V. Ground color very pale green (or the alternative pink or fawn) 

Sphinx chersis 

V. Ground color bright grass green (or crimson or brownj W 

W. Only six stripes are fully developed (pink); head broad and rough to the 

naked eye Dolba hylaeus 

W. With seven equally well-developed stripes X 

X. Horn green, black at the sides, straight; stripes with pink 

Sphinx gordius and luscltlosa 

X. Horn deep red (as also the stripes on the head); stripes violet 

Sphinx druplferarum 

X. Stripes heavily marked with black, and often shaded with blue 

Sphinx kalmiae 

Y. With oblique stripes running up toward the front Z 

Y. With pale transverse stripes; black Pseudosphinx tetrlo iFla.) 

Y. Catocala-like with small horn ami a tubercle on middle of abdomen 

Madoryx (M. pseudothyreus occurs in Fla.) 

Y. With subdorsal eyespots ... A 

Y. Checkered or with patches of yellow dots . . .c 

Y. Metathorax sharply humped with a dorsal eyespot E 

Y. More or less striped longitudinally . . .F 

Z. Oblique stripes and subdorsal red Proserpinus gaurae 

Z. Oblique stripes and subdorsal yellow Pachylia ficus (Fla.) 

A. A single eyespot on segment Al Xylophanes porcus ? (Fla.) 

A An eyespot on Al dark-pupilled, the rest light-pupilled .'... Xylophanes tersa 

A All the eyespots alike b 

B Ground color bright green; last eyespot quite like the others, lower part of 

head light Dellephila llneata 

n Ground color olive green; last eyespot often stretching out toward the horn; 

head with a black band below Deilphila gallli 

c Horn very slender Daremma catalpse 

c Horn normal D 

D Subdorsal yellow powdering if present continuous with that lower on the 

sides; lower part of face black Dellephila gallil 

D Patches of subdorsal yellow powdering on each segment; or checkerol with 

black and green; lower part of face concolorous Dellephila lineata 

E Eyespot black with a pale ring Erynnyis alope 

E Eyespot black, with some red behind Erynnyis ello 

E Eyespot red, with a black center Erynnyis cramerl 

F Horn slender; with red dorsal stripe and two white stripes or rows of white 

.spots on the sides Hyloicus cupressl 

F Horn various; otherwise marked, not feeding on pine G 

G Front edge of cervical shield raised and rough H 

G Cervical shield lightly and evenly granulated 

Younger stages of Proserpinus flavofasclata, Hemarls, and very 
young stages of many Sphingidae. 
H Dorsal dark stripe edged on both sides with pale; horn much longer than 

head Hemarls thysbe 

H Dorsal dark stripe vague; horn about as long as head Hemarls dlffinis 

1911] Study of Caterpillars of Sphiugidae 279 



Fig. 1. Front view of head of Sphinx liguslri. It is fairly typical of the spe- 
cies in which the head tapers moderately, but more rounded out on the sides than 
the average. The front is also wider and less lobed at the bottom. 

Fig. 2. Labrum of Hemaris Ihysbc. Compare Annals E. S. A. Ill; PI. xiv. 
Fig. 50. 

Fig. 3. Proserpinus proserpina. Typical of the SemanophorEe; compare also 
Figs. 6 and 14. 

Fig. 4. Front view of head of last stage of Ellema harrisii. The triangular 
head, which is more typical of the Smerinthinae. 

Fig. 5. Labrum of the same. Hyloicus is similar. 

Fig. 6. Labrum of Hip potion ceterio. 

Fig. 7. Labrum of Pachysphinx modesla. P. occidentalis .is the same, and the 
normal Smerinthina; differ only in having one less marginal seta. 

Fig. 8. Labrum of Sphin.\ (Lintneria) eremitus. typical of the lower 
species of Sphinx. For one of the higher type see Annals E. S. A. Ill, PI. xiii. 
Fig. 40, which is 5. gordius, labelled "Dolba hylaeus" in error. The species 
labelled gordius is certainly 5. drupiferarum. 

Fig. 9. Deilephila lineala. The other species are about the same. 


Fig. 10. Front view of head of Hyloicus pinastri. Sphinx drupiferarum is 
similar in outline, and so are all the species described as having a broadly rounded 

Fig. 11. Labrum of Dareinnia catalpae, showing seta iii high, as in the Semano- 

Fig. 12. Next to last stage of Cressonia juglandis. In the same stage Lapara 
has the same peculiar shape. 

Fig. 13. Labrum of Cressonia juglandis, last stage. 

Fig. 14. Labrum of Amphion nessus. Typicat 

Fig. 15. Head of Hippolion celerio, showing the slightly squarish form which 
is most frequent in the Semanophora:. 

Fig. 16. Head of Hemaris croatica. It is broader than our species of Hemaris, 
and resembles Macroglossa except in the small front. 


Vol-. IV, Plate XVII. 

n'iUiam T. M. Forbes. 

Annals E. S. A. 

Vol. IV, Plate XVIII. 

William T. M. Forhrs. 


By Miriam A. Palmer. 

It is the purpose of this paper to give a report of some 
experimental investigations in heredity which were made with 
four different forms belonging to the genus Adalia Mulsant. 
These forms shall herein be designated as melanopleura Leconte, 
annectans Crotch, coloradensis Casey, and humeralis Say.f 

ilfe/a»o/?/ez<;'a (Fig. A, PL XIX), as met with in this paper, is 
described as follows: Head black, with fine apical line of 
white, and with a whitish triangle next each eye with the apex 
pointing mesad and nearly reaching the median line. Some- 
times a median strip of whitish connects these spots and extend- 
ing forward to the apical line, which may also widen, leaves 
only a pair of black spots or brownish dots, one on either side of 
the median line on the anterior part of the head. (See head 
markings in Figures A, B, C, and F, PI. XIX). Pronotum pale, 
with black M-shaped design and a lateral black spot, except in 
an unusually albinic form where the spot is absent or repre- 
sented by a mere dusky area. The black spot when present 
may vary from a moderate sized area well enclosed by the 
surrounding white, to a large area which may break more or 
less widely through the surrounding white so as to connect 
with the M design. See pronotum markings in Figures A, B, 
and C, PI. XIX). The basal marking is usually large in this form 
but may be rather small in some cases. Elytra brownish red 
and immaculate, or with faint dot on lateral margin of each 
elytron. Legs yellowish brown, darker on outer margin. 
Length 4-6 mm., width 3-4 mm. 

Annectans, Figures B, C, and F, Plate XIX, includes quite a 
range of variation. The group as met with in this study is 
described as follows: Head as in melanopleura, pronotum as 

*This paper is an outgrowth of breeding cage work with the Coccinellids, 
assigned me by Professor Gillette as a part of his Adams Fund project on Life 
Histories of the Plant Lice and Their Enemies. 

t These determinations are according to Major Thos. L. Casey, who very 
kindly criticized my determinations of the forms referred to in this paper, excepting 
that annectans includes also an unusual and rather rare form (Fig. F, PL XIX), the 
status of which seems to me a little uncertain, but which Mr. C. W. Leng deter- 
mines as annectans. Lacking any biological proof to the contrary I have included 
it under annectans. 


284 Annals Entomological Society of America [Vol. IV^ 

given for melanopleura except that the basal marking is on the 
average somewhat smaller and is, in rare cases, even absent. 
The lateral black spot also is absent in an unusually albinic 
form. Figure F, Plate XIX. Elytra reddish yellow, usually lighter 
than melanopleura, quite yellowish for several weeks after 
emergence, becoming redder with age, though some never 
develop much of the red color. In the individuals reared of the 
more albinic form. Figure F, Plate XIX, the red color began to 
appear immediately after emergence but was paler in the region 
of the spots, giving a sort of blotchy appearance. This paler 
area may persist even in old beetles which have hibernated. 
Each elytron typically with a longitudinal posteriorly pointed 
black dash from the base at each side of the suture, and two 
sub-basal spots, the outer more basal, also with a transverse 
series of three black spots just before the middle, and two more 
at apical fourth, the outer very close to the margin. These 
black spots may vary from mere dots with some absent, to 
large blotches which may have more or less tendency to con- 
fluence; so that in color pattern many resembled ovipennis 
Casey, and a very few came very close to transversalis Casey 
as figured by Johnson.* Mr. Casey, however, to whom I sub- 
mitted specimens of these beetles, says that they are not his 
species as they do not show the proper punctuation. Some 
specimens show a rather definite pattern of red spots, two on 
each elytron, one a large oblong spot at the humerus and the 
other a smaller round spot close to the suture and between the 
middle and apical series of black spots. Legs and size as in 

In the more albinic form. Figure F, Plate XIX, the anterior 
spots w-ere always lacking and the middle and apical series 
were irregularly represented. Altogether this form differs from 
the rest of the group in three respects; namely, in lacking the 
anterior or basal elytral black spots, the absence of the lateral 
black spots on the pronotum, and in the presence of the redder 
coloration of the elytra. Three individuals of this form 
appeared in one batch of annectans, and one in another batch of 
amiecians, also two from a batch of larvae, from melanopleura 
parents, which produced both melanopleura and annectans. In 
these broods there were also some individuals which might be 

* Johnson, Roswell H., 1910 — Determinate Evolution in the Color Pattern 
of the Lady-beetles, Carnegie Institution of Washington. Pub. No. 122. 
Papers of the Station for Experimental Evolution, No. 15. 

1911] Heredity in Adalia ' 285 

considered as intergrading forms to some extent. This form 
presents a rather strikingly different appearance from the rest 
of the group, since all of these characters seem as a rule to go 
together, making a rather pronounced gap in the series of 
variation. It seems indeed to be closer to coloradensis than to 
annectans. Furthermore, among the 109 annectans which 
were reared from eggs of two annectans females and two melano- 
pleura-annectans hybrid females, mated with one annectans 
male and one melanopleura male, not one of these forms 
appeared, which fact seems to show that it is not a common 
fluctuating variation at least. It seems that the heredity 
might be segregate and experiments are now in progress to 
determine this point. It is on this account that I have thought 
best to call attention to it separately though for the rest of this 
paper it will be included under annectans. 

Coloradensis Casey, Fig. E, PI. XIX, is described as follows: 
Head black with fine apical margin of whitish, and triangular 
pale spot next each eye as found in annectans. Pronotum 
black with very fine apical pale margin sometimes obliterated, 
the posteriorly pointed median pale dash from the apical 
margin very small when present, sides with same pale pattern 
as melanopleura but lacking the black lateral spot, basal mark- 
ing absent. Elytra brownish red, about the same color as 
melanopleura, with a duplex black spot at the middle, some- 
times in the form of a band, sometimes appearing as two sep- 
arate spots, also a similar series of spots, two in number at 
apical fourth, the inner one the larger. Legs and size as in 
nielafw pleura, but the shape perhaps a little more narrowly oval. 

Humeralis, Say, Figure D, Plate XIX ; Head same as in melano- 
pleura. except that sometimes a different pattern appears as 
shown in Figure D, Plate XIX. Pronotum black with fine apical 
line and narrow side margins pale, apical line sometimes oblit- 
erated, basal marking always absent. Elytra black with a 
large oblong yellowish red to bright red spot at humerus and 
another small round one at three-fifths and close to the suture. 
The red marking on the elytra seems identical with the red 
pattern above mentioned as appearing in some specimens of 
annectans. Legs and sixe same as given for melanopleura, shape 
usually a little more rounded posteriorly. 

The work with these forms was at first undertaken merely 
for the purpose of obtaining specimens for life history drawings 
of melanopleura and annectans. In rearing these forms the 

286 Annals Entomological Society of America [Vol. IV, 

fact of their interbreeding with each other and with humeralis 
was discovered, and then the work was directed along the line 
of heredity investigation. About four hundred beetles were 
reared to maturity from about three thousand eggs hatched. 
These beetles proved much more difficult to rear than the 
larger species such as Hippodamia convergens and Cocci nella 
quinque-notata because of their more limited range of food and 
more delicate constitutions. All large aphids brought disaster 
in the breeding cage and sometimes even the small cottonwood 
louse, Chaitophoriis populicola Thos. was rejected. In the latter 
case it was perhaps due to an odor left by a certain species of 
attendant ants, since these lice did not always prove objection- 
able. One feed of unfavorable lice would sometimes cause the 
death of from one-half to nine-tenths of a cage of larvae. 

Work was begun with these beetles with the capture, 
May 13, 1910, in the foothills near Fort Collins, Colorado, of 
a pair of annectans. Eggs of this female were laid in the lab- 
oratory and the larvae reared. When the beetles eight in 
number emerged, four proved to be like the parents and four 
were humeralis. Three subsequent batches of larvae giving 
nine adults, were reared from eggs laid by this female and her 
spotted daughters with the result that three of the beetles 
were annectans and six were humeralis. 

For the purpose of obtaining more material and also of 
ascertaining how frequently such mixing occurred, two or three 
dozen pupae of this species were collected outdoors, and as soon 
as the beetles emerged and the colors developed the different 
forms were isolated in separate cages. About fifteen percent 
were humeralis and the rest were about evenly divided between 
melanopleiira and annectans. The humeralis beetles escaped 
by accident, but from the eggs of the other forms a considerable 
number of larvae were reared to maturity. From the eggs laid 
in the melanopleiira cage thirty beetles were reared, and in each 
batch a large proportion were annectans, sometimes over half 
the batch and once the entire batch. Practically the same 
proportions were obtained from eggs of one or two females 
captured at other times. Besides the forms already mentioned 
three individuals of coloradensis appeared among the progeny 
of the above mentioned cage. Unfortunately these were not 
used for breeding purposes but were pinned up and put in the 
collection. Breeding experiments are now, however, in prog- 
ress with this form. 

1911] Heredity in Adalia 287 

From the cage of annectans only annectans were obtained. 
Twenty-five adults were reared from eggs laid in this cage, and 
fifty-seven from eggs laid by a female tested in a way to be 
explained later, making eighty two beetles in all, and every one 
proved to be annectans. The eggs of one annectans female 
captured out of doors produced several melano pleura but this 
female had probably been fertilized by a hybrid male or even by 
both melanopleura and annectans males before it was captured. 

The hiimeralis beetles reared from the first pair mentioned 
were used for breeding purposes and all the individuals reared 
came true to type, about thirty beetles maturing. 

It was now indirectly evident that mixing was quite com- 
mon between melanopleura and annectans and that it sometimes 
occurred between annectans and hiimeralis, but there was no 
evidence that it occurred between melanopleura and humeralis. 
To ascertain whether this latter were possible and also to make 
the actual crosses in the other cases in order to further investi- 
gate the law of heredity, efi^orts were made to cross humeralis 
as often as possible with annectans and melanopleura. Humer- 
alis was found to hybridize just as freely with one form as with 
the other. No more difficulty was encountered than would be 
expected even among members of the same form under the 
same circumstances. On one occasion an annectans male chose 
a humeralis female even though a female of its own kind was 
present in the cage. 

Unfortunately only one female of humeralis was available 
for this purpose sufficiently early in the season, but there were 
several males which proved capable and these were crossed 
with females of both melanopleura and annectans. The female 
of humeralis that was used was probably the one that produced 
all of the above mentioned 30 humeralis, all true to type, she, 
at any rate produced a large proportion of them. This beetle 
was crossed with an annectans male but she died so soon that 
only two beetles were reared from this union. They were 
annectans but were too feeble for further breeding. An annec- 
tans-humeralis hybrid female was mated with a melanopleura 
male and later with an annectans male. This female had 
previously been kept in a cage with its brothers and the eggs 
laid had produced seven humeralis and four annectans, but 
after these crossings no more humeralis appeared though 
forty-seven beetles were reared. Three crosses were made by 
means of the humeralis males and melanopleura and annectans 

288 Annals Entomological Society of America [Vol. IV, 

females and from these 169 beetles were reared in the first 
generation. All but one were either melanopleiira or annectans 
according to the composition of the female. This one excep- 
tion was a humeralis beetle. A noticeable character of the 
progeny of these crosses was the greater vigor of the individuals 
so that a larger percent matured as compared with the purer 
strains. From one of the above three pairs, an annectans 
female and a humeralis male, the first generation of which con- 
sisted of fifty-seven annectans, four second generation beetles 
were reared and they proved to be two annectans and two 
humeralis. The beetles then refused to lay any more eggs and 
seemed to be preparing for hibernation. They had been 
unavoidably subjected for a few days to a temperature low 
enough to stiffen them up considerably and cause them to 
nearly cease eating and the subsequent removal of them to an 
almost summer temperature, though it caused the eggs to 
hatch in half the time they had under the low temperature and 
increased the appetites and rate of growth of the larvae quite 
remarkably, failed to cause the beetles to lay any more eggs. 
Work had, therefore, to close for the season at this interesting 
point, and the beetles were put into hibernation.* 

From these crosses there is another lesson to be learned 
besides the relation of annectans and melanopleura to humeralis. 
namely; something about the heredity between melanopleura 
and annectans themselves. The process of mating these forms 
with humeralis which is recessive to both, served as a test of the 
germinal composition of the member of the pair carrying the 
dominant characters. In the case where two melanopleura 
females, which had been isolated from annectans from time of 
emerging were crossed with humeralis males there were pro- 
duced 29 melanopleura to 25 annectans, and 31 melanopleura to 
26 annectans respectively. Melanopleura was in each case a 
little in excess of 50 percent. In the case of the annectans- 
humeralis hybrid female mated with the melanopleura-annec- 
tans male the progeny was 19 melanopleura and 28 annectans. 
The higher percent of annectans was doubtless due to the fact 
that an annectans male was put into the cage during the latter 

* Just as this article was ready to send to the publisher a lot of second genera- 
tion beetles, from the melanopleura females crossed with the humeralis males, 
emerged. From the eggs of the first generation melanoplenra-humeralis hybrids 
there were reared 19 melanopleura and 7 humeralis. From the eggs of the first 
generation annectans-humeralis hybrids there matured 12 annectans and 5 humeralis. 
These figures come very near to the Mendelian ratio for progeny of hybrids. 

1911] Heredity in Adalia 289 

part of the period, because just before the last three batches the 
proportion was 16 melanopleiira and IS annectans, and the last 
three batches gave 3 melanopleura and 10 annectans, thus mak- 
ing a sudden change in the proportion. This male was in ail 
probability pure annectans as there has not been found, in my 
experience, any proven case of annectans carrying melanopleura 
characters. The characters carried by the female could have 
had no influence whatever in the results, since neither of the 
characters carried by the female was dominant to the char- 
acters carried by the male. Either the melanopleura or the 
annectans characters of the male would realize themselves 
whether they met an annectans or a humeralis character of the 
female. These results approximately show that the melano- 
pleura-annectans hybrids carry the characters in the proportion 
of half and half. The somewhat high percentage of melano- 
pleura obtained in these cases was more than balanced by the 
extremely low percentage obtained in the case of the progeny of 
the cage of melanopleura-annectans hybrids, in which case 
melanopleura constituted less than half of the progeny when it 
should have constituted three-fourths. The mortality in this 
latter case, however, was so great that the data are hardly 

Another melanopleura female from melanopleura-annectans 
hybrid parents after being fertilized by some of its melano- 
pleura brothers was isolated for a few days, during which time 
it laid three batches of eggs. From these eggs were reared 
9 melanopleura, 3 humeralis, and 1 annectans. Excepting the 
one annectans, this was just the right proportion for the progeny 
of two hybrids according to the Mendelian law. This annectans 
individual, (if it did not get in by mistake which was very 
unlikely, great care having been exercised) must have been due 
to fertilization by a melanopleura-annectans male probably 
before the melanopleura-humeralis male. The female was 
then mated with a humeralis male and after that 14 adults 
were obtained, 6 melanopleura and 8 annectans. The results in 
this case seem to indicate that there had been a cross between 
the melanopleura ancestors of this female and humeralis, while 
still in nature and that in the first generation reared in captivity 
the dominant melanopleura had kept it concealed, so that it 
was not until the second generation that the crossing between 
two hybrids happened to take place, -thus allowing the humeralis 
character to appear. 


Annals Entomological Society of America [Vol. IV, 

From the cross between an annectans female (reared from 
melanopleura parents) and a humeralis male 57 beetles matured 
all annectans. This showed the female to be pure strain 
tl^iough descended from melanopleura parents. 

The foregoing results are given below in tabulated form: 

>2d gen. 


1 Male — annectans 1, . 
Female — humeralis j ° 

2 Male — humeralis Ust gen. 

Female — annectans |2nd gen 

3 Male— humeralis j, . 

Female — melanopleura *( ^ ' 

4 Male — humeralis 'i , . 

Female — melanopleura*! 

5 Male — melanopleura* , 

and later annectans list gen. < 

Female — annectans-humeralis ' 


Humeralis Hybrid 

Female — annectans-humeralis . \ a ■ 
Male — annectans-humeralis....] " 8 ■■■ 

Female — melanopleura-humeralis 1 

Male — melanopleura-humeralis (probably). 
Also melanopleura-annectans (prob.) .J 
Female — melanopleura-humeralis \ a 
Unle-humeralis joffsprmg. 

Melanopleura-annectans Hybrids 

Cage of males and females — offspring 

Female captured, male annectans? 

— offspring 
Two females and one male ] 

mated with recessive. . . . ^off spring. 
(See crosses 3, 4 and 5) j 


Cage of males and females — offspring 

One female, captured, male unknown, 


One female with humeralis male (See 

cross 2) — offspring 


One female with 3 males — offspring 

(Female used later in cross 1) 


* melanopleura-annectans hybrid. 





















































1911] Heredity in Adalia 291 

From the foregoing results the following conclusions seem 
to be quite evident: 

I. Melanopleura is dominant over aniiectans, coloradensis, and humeralis, 
and the heredity is segregate. 

1. Over annectans since 

a. The hybrid fonn between melanopleura and annectans is 

melanopleura. Of the progeny, 30 in number, of a cage of 
melanopleura, anneclans constituted over half. In the 
progeny of two females and one male tested by mating 
with humeralis there appeared 76 annectans and 69 
melanopleura, altogether, which is very close to the 
Mendelian ratio for the segregation of characters in 

b. Annectans has in no case given evidence of carrying melano- 

pleura characters. The 23 progeny from a cage of 
annectans showed no melanopleura characters nor did any 
of the 57 progeny of the annectans female mated with 

2. Over coloradensis since the hybrid fonn between melanopleura 

and coloradensis is melanopleura as is shown by the fact that 
3 coloradensis appeared among the offspring of melanopleura 

3. Over liumeralis since 

a. The hybrid fonn between melanopleura and humeralis is 

melanopleura. In "the first generation from three crosses 
of melanopleura with humeralis or with annectans-humeralis 
hybrids, humeralis appeared but once among 159 individ- 
uals. A melanopleura-humeralis female mated with its 
brothers gave 9 melanopleura, 1 annectans, and 3 humeralis. 
The same female mated with a humeralis male gave 6 
melanopleura and 8 humeralis, approximately showing 
the segregation of characters to be according to the 
Mendelian law. The second generation from crossings of 
melanopleura with humeralis consisted of 19 melanopleura 
and 7 humeralis. 

b. Humeralis has given no evidence of carrying melanopleura 

characters. The 30 offspring from humeralis parents all 
came true to type. 
II. Annectans is dominant over humeralis since 

a. The hybrid fonn between annectans and humeralis is 

annectans. In the cross between annectans and humeralis 
humeralis did not appear at all in the first generation of 
57 progeny, but did appear in half of the second gen- 
eration which consisted of 4 beetles. Annectans-humeralis 
hybrids mated with each other produced 7 annectans and 
10 iiumeralis in one case, and in another 12 annectans and 
5 humeralis. 

b. Humeralis has given no evidence of carrjdng annectans 

characters, as shown by the 30 offspring of humeralis 
parents all true to type. 

292 Annals Entomological Society of America [Vol. IV, 

This subject is still unfinished and experiments are now in 
progress to determine the relation of coloradensis and the 
rather albinic form of anneclans to the other forms. 

It would be interesting to interbreed these forms with other 
species of Adalia, especially with the European /r/g/Ja Schneider 
and with bipunctata Linneaus. 

Observations were also made on the beetles used in the 
foregoing experiments for the purpose of ascertaining the 
heritabilit}' of the characters of the spots on the elytra in 
annectans and of the markings of the pronotum in this same 
form and in mclanopleura. The progeny resulting from the 
mating of annectans and melanopleura beetles with the recessive 
hnmeralis were examined when the number was large enough 
to afford sufficient data. The beetles in these cases were par- 
ticularly advantageous for this purpose because the dominant 
characters would be the only ones to show in the first genera- 
tion, thus reducing the number of strains which would appear 
to one or two. In the case of the melanopleura-ajinectans 
hybrids there would be one strain of annectans and one of 
melanopleura, which would afford a very simple series and show 
very plainly whether these characters behave at all as unit 
characters or whether they seem to be fluctuating variations. 
The results are shown in the drawings Figures 2 to 7, Plates 
XX to XXII. 

In the markings of the pronotum, special attention was paid 
to the character of the lateral black spot and the extent to 
which was it enclosed by the surrounding white. The median 
posteriorly pointed dash of white from the apical margin and 
also the basal marking of whitish are sometimes very small or 
even absent; but in this study only secondary attention was 
paid to these and the drawings, except curve (e). Fig. 7, Plate 
XXII, are arranged in series according to the aforesaid black spot. 
The pronota of melanopleura and annectans are arranged 
separately in each case. 

In the case of the elytra primary attention was paid to the 
confluence of the spots, and the series is arranged according to 
the number of confluences in each case. The parents of each 
series are drawn in full or designated above and the first gen- 
eration progeny in a row below. The numerals below each 
drawing indicate the number of individuals in that class. As 
the hnmeralis parent seems to have no influence on the char- 

11)11] Heredity in Adalia 293 

acters of the first generation it was not thought necessary to 
draw this parent. 

Figure 2, Plate XX represents the aiinectans-hmneralis hybrid 
female and her progeny resulting from union with a melano- 
pleiira-annectans hybrid male, and also for the last few days of 
the experiment, with a pure annectans male. The numbering 
of the spots is after Weise taken from Johnson 1910. In this 
case the progeny would contain four strains of annectans, one 
from the mother, one from the nielanopleura-annectans father, 
and two strains from the annectans father, which, however, 
could hardly have affected more than the last three batches of 
eggs. This would be just the number of strains to be repre- 
sented if two members of annectans were mated. The males in 
this case were both lost and so can not be shown in the figure. 
Of the batches after the annectans male was introduced, in the 
elytra series, one beetle was in class (d), six in class (e), 
three in class (f), and one in class (i). In the pronota 
series four were in class (k) and seven in class (1). There was 
considerably less variation among these than in the foregoing 
batches, but whether it was due to the annectans male or to 
environmental influences can not be ascertained with certainty; 
but as these were reared later in the season than the foregoing 
batches, during the latter part of August and the early part of 
September, during which time an unusually cold wave occurred, 
the only environmental influence would probably have been a 
lower temperature. This factor, however, would, from the 
experience of Tower* and Johnson, be expected to produce a 
melanic effect, but here the difference was albinic rather than 
melanic, so the case does not seem to be explained by the 
environmental factor, and unless it was produced by some 
unknown cause, seems most probably to have been due to 
heredity factors introduced by the annectans male. 

It will be noted in this case. Figure 6, curve (a), Plate XXI, 
and Figure 7, curves (a), and (b) Plate XXII, that all of the 
beetles, of both melanopleura and annectans, which were reared 
from this female were rather at the albinic end of the scale as to 
both elytral and pronotal characters. In the elytra none 
have more than two full confluences and the mother ranks at 

* Tower, William Lawrence, 1906. An Investigation of Evolution in Chrysom- 
elid Beetles of the Genus Leptinotarsa. Carnegie Institution of Washington, 
Pub. No. 48. 

294 Annals Entomological Society of America [Vol. IV, 

about the middle of the series and at one of the highest points 
of the curve. In the case of the pronotum the mother was 
decidedly more melanic than the apex of the curve for either 
annectans or melanopleura. The curves for these two forms 
were not alike, annectans having the greatest number, 45 per 
cent at the albinic end of the scale with the black spot well 
enclosed by the surrounding white. In the melanopleura 
series only 11 per cent were at this point, the largest number, 
83 per cent, having the black spot rather weakly enclosed. 
None of the annectans here showed the red pattern on the 
elytra, as shown in Figure B, Plate XIX, though the mother 
shows it faintly. 

Figure 3, Plate XX, represents the annectans female crossed 
with humeralis male. In this case we would expect to find 
only two strains of annectans. Here, however, the variation 
was considerably broader than in the former case where four 
strains were represented, the curve beginning at the same 
point of albinism as the former case and extending to four and 
a half confluences (that is to four and a pronounced tendency 
to a fifth confluence), Figure 6, curve (b). The mother was 
several degrees more albinic than the highest point of the 
curve. Note here that in the mother there is an absence of 
spot 4 and also that there is a small spot between spots 1 and 2, 
which, though very unusual, probably denotes tendency to 
confluence between spots 1 and 2. Neither the presence of 
this extra spot nor the absence of spot 4 show in any of the 
progeny examined, though both confluence and tendency' to 
confluence appear between spots 1 and 2. The mother of these 
seems to have shown nothing of the red pattern mentioned 
above and shown in Figure B, Plate XIX, but in the 37 off- 
spring, 9 showed it very plainly, 16 moderately plain, 4 fainth-, 
and in 6 it was absent. 

Figure 4, Plate XXI shows a melanopleura-annectans hybrid 
female, crossed with a humeralis male, and her first generation 
progeny. Here there can be but one strain of annectans to 
appear in the progeny. The curve of variation. Figure G, curve 
(c), Plate XXI, covers a somewhat wider range of variation 
than in the case of the first instance, curve (a) where four 
strains are represented, the largest number of confluences being 
three. Here 23 out of 27 or 85 percent lack spot 6. In the pro- 
nota of annectans a peculiarity was observed in that sometimes 
either the basal marking or the apical median dash were lacking. 

1911] Heredity in Adalia 295 

For these pronota two curves were given, Figure 7, curves (d) 
and (e), curve (d) to show the variation of the lateral spot only 
and (e) to represent the general melanism when the other 
markings are considered, each degree representing about the 
equivalent of the melanism of the state of the lateral spot as 
given in the legend for the respective columns. The curve for 
annectans in this series was much broader than that for melano- 
pleura. The mother was rather toward the albinic end of the 
series for annectans and at the melanic end for melanopleura. 
All of the annectans, 27 in number, had the red spots on the 
elytra, as shown in Fig. B, Plate XIX. 

Figure 5, Plate III, shows another melanopleura-annectans 
hybrid, female mated with a humeralis male, and her first 
generation progeny. Here again would be but one strain of 
annectans. The range of variation in the elytra of annectans 
was not very broad, showing none of the more albinic forms, 
the curve. Figure 6, curve (d), Plate XXI, beginning at one con- 
fluence and extending to three and a half confluences. In the 
pronota of the annectans series, Figure 7, curve (g), Plate XXII, 
uniformity almost obtains, 93 per cent having the lateral spot 
well enclosed and 7 per cent being one-fifth enclosed. In the 
melanopleura series, however, the curve. Figure 7, curve (h), 
Plate XXII, is very broad extending to a degree of melanism that 
is quite rare. The mother ranks at the albinic end of the scale 
though the highest part of the curve for her melanopleura 
offspring is four degrees further to the melanic end of the scale. 


In comparing the curves for the elytra it must be born in 
mind that the chief points of comparison are the melanic 
positions of the range and highest points of the curves. Since 
the number of individuals represented by each of the curves 
was not uniform, the exact number on any one line shown by 
the different curves is not truly comparable; only the melanic 
position of high and low points and range in each curve can be 
compared with the same in another curve. 

It will be noted that each curve has one or two points that 
are much higher than any other points in the curve, and that 
these high points in the different curves vary greatly in melanic 
position, also that the curves vary considerably in their range. 
It seems as though these high points in the curves might repre- 
sent centers of variation. The curves would then signify that 
different strains of these beetles have different centers of 

296 Annals Entomological Society of America [Vol. IV, 

variation and different scopes of variation. Curve (c), Figure 6, 
Plate XXI, which represents but one strain of annectans covers a 
wider range than curve (a) which represents four strains. 
Curve (d), which also represents one strain is quite narrow, 
seeming to signify that this strain had a greater degree of con- 
stancy than the others. The mother in each of these cases 
occurred within the range of variation for her progeny but not 
always at the highest point of the curve though in both of the 
instances where this observation was possible she occurred at 
one of the high points, see curves (a) and (b), Fig. 6, Plate XXI. 
Two of the mothers being melanopleura had no place in the 
elytra series, and since in the cases where the mother was 
annectans two or more strains were represented, the fact of the 
highest part of the curve not being at the same position as the 
mother might in this case be explained as due to one of the 
other strains involved. 

There seemed in some cases to be a certain measure of her- 
itability of different characters in the color pattern of the 
elytra. The absence of spot G in Figure 4, Plate XXI, seemed to 
be inherited to a large degree since it was lacking in 21 out of 27 
beetles. The mother being melanopleura could not be observed 
on this point. This spot seems from my observations to be the 
one most frequently lacking in this form, indeed almost the 
only one except in a small minority of beetles. Spot 4 was 
absent in only three beetles in this study, in Figure 3 (a), 
Plate XX, and in two others not drawn but ranked with (f) and 
(g) respectively in Figure 2, Plate XX, spot 5 was faint in one. 
Figure 3 (c). The absence of spot 4 seemed not to be inherited 
in these cases, as no case of absence occurred in the 37 progeny 
of the mother, Figure 3 (a), which lacked it, and it appeared 
only twice in the 30 offspring in Figure 2, Plate XX. 

Some observations were made on the order in which con- 
fluences take place. Spots 6 and 7 seemed to be the first as a 
rule to connect, as in this study there was only one instance 
where a beetle showed confluences and had these spots separate, 
see Figure 3 (e) . There were three such cases where spot 6 was 
absent, but even in the case of absence there was often a pro- 
jection toward its position from spot 7 as though in these 
instances the confluence was even more persistent than the 
spot itself. After this confluence no further order was observed 
except that between spots 4 and 5 it seemed to be the most 
unusual and perhaps the last in order. 

U)ll] Heredity in Adalia 297 

In the case of the pattern of reddish spots on the elytra of 
a nnectans Figure B , Plate XIX , it seemed as though there might be 
segregation in some cases at least, and that the absence of the 
character was dominant to its presence. In the series in Figure 
2, Plate XX, it shows faintly in the mother (the dimness may be 
due to fading after death as this character was not recorded 
during life) and it was plainly evident in the mother and a 
brother of this beetle, in fact in all of the individuals of this strain 
that have been preserved. It shows in none of the 30 progeny 
of this beetle, but this absence -may be explained as due to 
the males, which being lost, can not be examined as to their 
possession of the character. 

In the series in Figure 3 where the mother does not show the 
marking but carries two strains of annectans, it appeared in 
five-sixths of the beetles to a greater or less degree. As the 
male in this case was humeralis both of these strains must have 
come from the mother and its absence in her development 
would seem to signify the dominance of the absence of the 
character over its presence. The proportion, however, found 
in the progeny seems rather puzzling unless the humeralis 
character from the male could have had any influence in the 
proportion, which seems unlikely. 

In the series in Figure 4, Plate XXI, it appeared in all of the 
27 annectans progeny. The mother, being melanopleura of 
course does not show it. In the series in Figure 5, Plate XXI, 
some show it and some do not. The exact number in each case 
can not be determined as some of the beetles have developed so 
much of the red color in their elytra during hibernation that it 
is impossible to tell with certainty whether they possess the 
character or not. The mother being melanopleura of course 
does not show the character. The fact that some clearly show 
its presence and others just as clearly show its absence when 
they are all from one strain of annectans seems to be evidence 
against segregation in this case. 

In the pronota curves in Figure 7, Plate XXII, the matter is a 
little more complex as there are both annectans and melanopleura 
to be represented for each female except one, Figure 2, Plate XX. 
As the curves for these two forms even when from the same 
parents were different in every case not only in the position of 
the apex but also in range and sometimes very different, it 
would seem that each strain keeps distinct ; that is, the pronotal 
•characters of annectans do not mix with those of melanopleura. 


Annals Entomological Society of America [Vol. IV, 

When, however, the characters of the mother are compared 
with those of her offspring which are of the same form as herself 
Httle uniformity was found. In no case did she rank at the 
highest point of the curve, neither did she ever occur at the 
lowest point, nor ever outside of the range of variation for the 
offspring. There seemed to be some degree of heredity but it 
was not constant. The results appear a good deal the same as 
in the elytra, that there are centers of variation and a certain 
limit of range that were inherited to a greater or less degree, but 
with no evidence of segregation of unit characters such as occurs 
between vielano pleura, annectans, and hunieralis. 


Since sending the foregoing article to the publisher results 
have been obtained in the experiments concerning the relation 
of coloradensis, the so-called albinic form of annectans, and a 
similar form of melano pleura to the other forms treated. The 
albinic form of annectans, so-called for want of a better name is 
above described separately under annectans and figured at F, 
Plate I. The albinic form of melanopleura is identical with 
that of annectans in pronotal characters, namely, it lacks the 
lateral dot, the lateral margin of the pronotum being broadly 
pale as in coloradensis; in all other characters it agrees with 
melanopleura. The results obtained are tabulated as follows: 




















al. A. 

al. A. 


al. M 


MandA ,\ 

g ' 

C and H 
Cand A 
Cand A 


C andM 


al. A 

al. M 

al. A. 

al. M. 


C and A 1 

C and M 
M and H 
M and H 
C and A 

C and M 

































*A means annectans; C, coloradensis; H, hunieralis; al. A. albinic annectans; 
al. M, albinic melanopleura. 

t These are given as shown by the offspring when not known from pedigree 

tThis male was, judging from appearance, an intergrade between annectans 
and albinic annectans. It lacked the basal spots on the elytra but possessed the 
lateral dot on the pronotum, which latter seems to be the ultimate distinguish- 
ing character. 

1911] Heredity in Adalia 299 

These results seem to furnish conclusive evidence 

1. That coloradensis is a good variety or type equal with melano pleura, 

annectans, and humeralis, acting as a unit character in heredity. 

2. That when crossed with annectans, coloradensis produces a blended 

hybrid, in both elytral and pronotal characters, namely the form 
above referred to as an albinic form of annectans. 

'S. That when crossed with melanopleura a blend is produced in the 
pronotal characters, identical with the annectans blend; but in 
the elytra melanopleura dominates entirely.* This form was 
mentioned in the description of melanopleura as a "more albinic 

4. That when crossed with humeralis coloradensis dominates perfectly 
so that the hybrid form is indistinguishable from the pure strain 

of coloradensis. 

It seems that in every instance the more albinic character 
dominated over the more melanic one ; for example : immaculate 
elytra, in melanopleura, dominate over the spotted ones of each 
of the other forms. The absence of the black lateral dot in the 
pronotum, in coloradensis, dominates over its presence in each of 
the other forms. The presence of the whitish basal marking on 
the pronotum, of annectans and melanopleura, dominates over its 
absence in coloradensis and humeralis. The absence of the 
basal elytral spots, in coloradensis, dominates over its presence 
in annectans. The usual absence of confluence in the median 
and apical series of spots in annectans dominates over the con- 
fluence in these spots in coloradensis. Humeralis which pre- 
sents the most melanic characters in every particular in both 
elytra and pronotum is perfectly recessive to each of the other 

The inheritance of the faint lateral dot on the elytra in 
melanopleura was observed in the specimens at hand but no 
law was ascertained. It seems to be a mere fluctuating 

The single humeralis beetle mentioned in the article as 
appearing among the first generation offspring in the 4th cross 
in the table, between humeralis and melanopleura parents, was 
tested in breeding. It proved to be a male and was put into a 
cage with two humeralis females, from the eggs of which seven 
progeny were reared to maturity. All of these were humeralis 

* This statement is to be reconciled with the statement in the foregoing ar- 
ticle that melanopleura is dominant over coloradensis by the fact that at that time 
the hybrid was considered as only a variant of melanopleura. 

300 Annals Entomological Society of America [Vol. IV, 

which seems to prove that the beetle in question was pure 
strain. This beetle may possibly have gotten into the cage by 
mistake in spite of the great care exercised as several dozen 
cages containing larvae of all the forms were being tended and 
cleaned daily. 

Five other offspring were reared from humeral is beetles 
obtained in these experiments, and these all came true, making 
42 progeny in all reared from humeralis parents, breeding true 
in every instance. 

Two humeralis beetles without dorsal spots were obtained 
as the progeny resulting from a cross between a melanopleura 
male, (evidently a humeralis hybrid) from out of doors, with an 
annectans-humeralis female representing the third generation 
of humeralis reared in the laboratory. All of the ancestors and 
progeny, two in number, of this female, by a former mating 
showed the dorsal spots normally developed. These two beetles 
were the only progeny obtained from this union and efforts to 
rear offspring from them, though they proved to be male and 
female, have thus far been fruitless, seemingly due to a weak 
constitution as the eggs hatch poorly. The male seemed 
weak and both beetles died soon. It would seem from this case 
that the absence of these spots dominated over its presence, 
which is contrary to the behavior of heredity with regard to the 
other characters of this group. If this is not the case the 
strain in the laboratory must have carried this character of 
absence through three generations without it having a chance 
to meet its equal so as to be able to realize itself. 

Another cross which was made between an annectans male 
and a humeralis female last August but which laid no eggs 
until this, the following spring, produced in the first geneartion 
26 beetles, all annectans. The humeralis female was later used 
in the first cross represented in the first table in addenda. 

1911] Heredity in Adalia 301 

Fig. a. Adalia melanopleura Lcconte. 
Fig. B. Adalia atiiieclatis Crotch. 
Fig. C. Adalia annectans Crotch. 
Fig. D. Adalia humeralis Say. 
Fig. E. Adalia coloradensis Casey. 
Fig. F. Adalia annectans Crotch. 
Fig. G. Adalia melanopleura (more albinic form). 

Fig. H. Pupa of .1. annectans, melanopleura, coloradensis, ami humeralis. 
Fig. \. Larva of .1. annectans, melanopleura, coloradensis and humeralis. 
Fig. J. Eggs of A. annectans, melanopleura, coloradensis and humeralis. 
All drawings are magnified o diameters. 

Fig. 1 shows in diagram the results of the foregoing experiments in inheritance 
between annectans, melanopleura, coloradensis, and humeralis. The numerals be- 
neath the circles in each case indicate the numlier of individuals in that class. The 
lines connecting with higher circles indicate parentage in each case. 

Fig. 2. a and b — characters of annectans-humeralis female, mated with males 
c to j — elytral characters of annectans offspring. 

k to n — pronotal characters of melanopleura and annectans offspring. 
Numerals indicate number of individuals in the class in each 
Fig. 3. a — elytral characters of annectans mother, mated with male indi- 
b to — elytral characters of annectans offspring. 
p to V — pronotal characters of annectans offspring. 
Numerals indicate number of individuals in each class. 

Fig. 4. a and b — characters of melanopleura annectans mother mated with 
male indicated. 
c to 1 — elytral characters of annectans offspring. 
m to / — protonal characters of annectans offspring. 
M to !' — pronotal characters of melanopleura offspring. 
Numerals indicate number of individuals in each class. 
Fig. 5. a and b — characters of melanopleura-annectans mother, mated with 
male indicated, 
c to d — pronotal characters of annectans offspring, 
e to k — elytral characters of annectans offspring. 
/ to r — pronotal characters of melanopleura offspring. 
Numerals indicate number of individuals in each class. 
Fig. 0. Shows the curves representing the variation in the elytral characters 
of the annectans offspring, drawn in the foregoing figures. The numerals on the 
left show the number of individuals. The degrees of melanism are designated by 
the legend below in each case, the most albinic being at the extreme left and the 
most melanic at the right. "Confluence )-'2"means one case of tendency to con- 
fluence, "confluence |" means two cases of tendency to confluence. The latter 
is here given a rank of its own as it does not seem equal in melanism to one full con- 
fluence. The curve above the legend "spot 6 absent" does not represent all the 
individuals lacking that spot, but only those with no case of confluence. 

Curve (a) represents the series in Fig. 2, Plate II. The full line triangle shows 
the position of the mother in this series. 

Curve (b) represents the series in Fig. 3, Plate II. The broken line triangle 
represents the position of the mother. 

Curve (c) represents the series shown in Fig. 4, Plate III. 
Curve (d) represents the series shown in Fig. 5, Plate III. 
The mother of the series for curves (c) and (d) were melanopleura and so have 
no place in this diagram. 

302 Annals Entomological Society of America [Vol. IV, 


Fig. 7 shows the curves for the pronotal series. 

Curve (a) represents pronotal characters of anneclans offspring shown in Fig. 
2, Plate II. 

Curve (b) represents pronotal characters of melanopleura offspring in Fig. 2, 
Plate II. 

The open triangle represents the mother of the series represented by curves 
(a) and (b), Fig. 2. a and b, Plate II. 

Curve c) represents the anneclans series in Fig. 3, Plate II. 

Curve (d) represents the anneclans series in Fig. 4, Plate III. 

Curve (e) represents the anneclans series in Fig. 4, Plate III, according to 
general melanism. 

Curve (f) represents the melanopleura series in Fig. 4, Plate III. 

The open triangle represents the mother, Fig. 4, a and b, Plate III. 

Curve (g) represents the anneclans series, Fig. 5, Plate III. 
Curve (h) represents the melanopleura series, Fig. 5, Plate III. 

The solid triangle represents the mother Fig. 5 (a) and (b), Plate III. 


Vol. IV. Plate XIX. 

Miriam A. Palm- r. 

Annals E. S. A. 

VOL. IV, Plate XX. 

y^eUn^pUura, cnneciar^s>^sU ha^e^c^s ■ 

0P r:^^ G)d (^P 




ann.ecta,ns hybrid O 
a.^ne ctans o 

e g 


1 =.30 

*:^ ^ m^ (^ 

annectans 1^ 

TrL^lcur\.ople UTap 
Fiq J 

b C d 


2 37 

,,J tThumeralii 6 

r g h i J H- I rn. n. o 

J 7 lis 24- 10 3 3 113 1'37 



f ^9 

Miriam A. Palmer. 

Annai;8 E. S. a. 

Vol. IV, Plate XXI. 

6 ^^3^ 
^^^^^ x-hu me rails cT 

• ••• 

11 Z 1 S 1 2 

anr^ ectans 


1 1 ' Z.S 20 10-30 

^jyK#/ X kumeralit, o 

anrtec ta.ns 

f g hi J_^ fC 

"^ -(^ f^ ff\ 

14 1 


z J -IS 


Fiq € 




























1 1 



. 1 





















f M 




d * 








■V' - 







%. -- t, ^ ^ ^ ^ ^ ^ ^ ^ ^ ^ ^ 

ne confluence 

a. ^5 2 ^Tnother of a, <^ Fig ^ 

b F^y 5 ^"j, , .6 <i ■■ S 

Miriam A. Palmer. 

Annals E. S. A. 

Vol. IV, Plate XXII. 






























\ ' 





\ ' 

Li 1 



\ I / 














-I \l 


\ \ 







1 ..i 

^-. \ 





' ^ 








\ > 











\ \ 




', \ 


^*> \ 












/\ maker 'i 

curve a, 1st. 9^^\ 

■„ b 

c . . . 

/\ rriother 
curue d 1st aen. 

▲ rnotKer 

curue q Ist'qeru 

h. .. 


:5? ^ 4? * * 

Minam A. Palmer. 


By P. E. Smith, 
Entomological Laboratory of Cornell University. 


The purpose of this investigation was to determine the 
value of the specific characters used in the descriptions of 
species of Pseudococcus. While all characters used have 
probably not been noted and while some of those omitted may 
be of considerable value still the more common and important 
characters have received attention. 

The investigation has been limited to five species,* namely: 
agrifoliae Essig, citri Risso, crawii Coq., longispinus Targ., and 
obscurus Essig. A large number of individuals in each species 
has been used giving a comparative study for specific variation. 

The writer wishes to thank Professor Alex. D. MacGillivray 
for the many invaluable suggestions given. 


An examination of the descriptions of species in this genus 
shows a great similarity in the characters used. Provided that 
these characters do not vary beyond certain limits, this would 
make the identification of the species easier. However, if these 
characters vary to any great extent and overlap and merge 
into each other, the adherence by systematists to these char- 
acters instead of the introduction of new ones is unfortunate, 
and would make the identification of species very difficult, if 
not impossible. In that case the most valuable data in the 
descriptions would be the locality and host-plant data. 

Body. 1. Size. -In the great majority of descriptions, the 
length and width of the body is given. In the measurements 
given, a great variation in the length of each species is noted.f 
Lengths such as 3-4 mm., 2-5 mm., 1.5-4 mm., are very fre- 
quent, showing the wide variation noted by those describing 
species. The tablesj showing the lengths of the body, (Tables 
1, 2, 3, 4, 5) in the five species studied do not show a variation 
any greater than this. A variation of from one and one-half 

* The writer does not express an opinion upon the validity of these species, 
t R. Mathewson. Can. Entom. XXXIX, p. 286. 

I In these and the following tables an ocular micrometer with a 2-5 and 1-8 
inch objective were used. All measurements are given in microns. 



Annals Entomological Society of America [Vol. IV, 

to twice the length of the smaller specimens is shown. A 
greater variation would probably be found if a larger number 
of specimens were measured. 

TABLE 1. PsEUDOcoccus obscurus Essig. 


Length of 

Length of 

Length of Setae 

Length of Setae 



of Anal-lobes 

of Anal-ring 



























































Length of 

Length of 

Length of Setae 

Length of Setae 



of Anal-lobes 

of Anal-ring 
















































Length of 

Length of 

Length of Setae 

Length of Setae 



of Anal-lobes 

of Anal-ring 











163 • 
































IX . 












Specific Characters in Pseudococcus 


TABLE 4. Pseudococcus crawii Coq. 


Length of 

Length of 

Length of Setae 

Length of Setae 



of Anal-lobes 

of Anal-ring 




















































TABLE 5. Pseudococcus longispinus Targ. 


Length of 

Length of 

Length of Setae 

Length of Setae 



of Anal-lobes 

of Anal-ring 



















































Also only the larger individuals were studied because of the 
danger of including those in the nymphal stage. Each species 
was collected from the same or neighboring host plants and 
were apparently under similar conditions, so that size cannot 
be taken as a specific character of any great importance. These 
measurements were taken from mounts. If unmounted spec- 
imens were used, the variation would probably be still greater, 
as then a second factor, namely, the amount of secretion pres- 
ent, would enter. Because of the great variation in size of the 
different individuals in a species the size of a specimen can be 
of but little if any value in specific determination. 

2. Shape. — In descriptions the shape of the body is vari- 
ously given as elongate-ovate, rounded-oval, convex, tapering 
at ends, elongate, etc., words which are nearly synonymous. 
A few species are stated to be considerably out of the ordinary 

312 Annals Entomological Society of America [Vol. IV, 

in shape, but in general synonyms are used in the descriptions 
of body shape. In the species studied, this could not be an 
important specific character as all the species studied were of 
practically the same shape. However, in some species, this 
might be a distinguishing character. 

3. Color. — In descriptions the color of specimens is usually 
given, being stated as, whitish, greyish, pinkish, with a red 
tinge, etc. The dermis of specimens in this genus is red. The 
color of the insect depends on the extent that this dermis is 
hidden by the waxy secretion. The amount of waxy secretion 
depends largely on the position in which the individual develops. 
Those in exposed position requiring more of the waxy covering 
as_ a jDrotection than those developing in well protected situa- 
tions. This variation in color is well shown in agrifoliae Essig. 
The color in this species is usually of a reddish, pinkish, or pink- 
ish brown. The writer has found specimens, however, ovipositing 
in exposed positions which were completely covered with the 
waxy secretion and through which the red dermis was not 
visible. Thus all variations from a white to a red specimen 
were found. In the other species studied the color was nor- 
mally white. In the five species studied color as a character 
for separating the species could not be relied upon. 

4. Segmentation. — In many descriptions the segmentation 
is referred to as distinct or not distinct. This is a character 
that is not used to any great extent. This is fortunate as the 
value of the prominence of the segmentation as a specific 
character is extremely doubtful. In mounted specimens it 
would depend to a large extent upon the method used in prep- 
aration. The segmentation in chloroform mounts is well pre- 
served. In caustic potash mounts it would be largely oblit- 
erated. In unmounted as well as mounted specimens the age 
and size of the specimens is of great importance. In nymphs 
of Pseudococcus the segmentation is very distinct. As it 
passes through its last moult and becomes an adult, this dis- 
tinctness of the segmentation is lost to a degree. Then as the 
insect becomes more and more distended with eggs this ten- 
dency to lose its prominent segmentation is increased. Also 
the distinctness of the segmentation in unmounted specimens 
would depend somewhat upon the amount of secretion covering 
the insect. With these factors it does not seem that the dis- 
tinctness of the segmentation can be a very satisfactory 
specific character. 

1911] Specific Characters in Pseudococcus 313 

Antennae. — The number of segments of the antennae 
is a generic and not a specific character, and so is of no import- 
ance in separating species. Several species have been described 
with seven segmented antennae. The number of segments, 
eight, of the type of the genus, longispinus Targ, has been 
adopted in this paper. However, a dimorphism has been 
described in two instances, viz. : by Folsom for trifolii Forbes, 
and by Essig for agrifoliae Essig. In these descriptions there 
is said to be a winter form which has seven segmented antennae ; 
this form giving rise to the summer form with the normal eight 
segmented antennae. If further investigation shows these 
observations to be correct, it will be of both specific and generic 

The comparative length of the different segments of the 
antennae is a character that is the most used of any of the 
characters of the insect's body. The relative length of the 
segments and the formulae deduced from the measurements is 
almost invariably contained in descriptions. Sometimes a 
considerable variation is noted, several formulae being given. 

In this study ten specimens of each species were used. 
Aside from the question of variation which will be taken up 
later, the relative length of the antennal segments is not a desir- 
able character to use. The greatest difficulty in its use is the 
difficulty of making correct measurements and the determina- 
tion of the exact limits of the various segments. The chitin is 
not continuous from one segment to the next and consequently 
the portion between the chitinous parts of the segments, the 
■conjunctiva, is not visible or only slightly so in well cleared 
specimens. Consequently, in making measurements, the deter- 
mination of the end of a segment will be only approximately at 
the center of the conjunctiva. This difficulty will be much 
increased if there are some bends in the antennae. A second 
difficulty is to determine some point at the end of the segment 
from which the measurement will always be taken. This 
difficulty is most apparent with the first segment. This seg- 
ment is an irregular truncated cone with the sides of different 
lengths, and but little longer than the width across the base. 
It will be seen that the determination of the same points for 
the measurement of this segment would be nearly impossible. 
In the tables given the writer does not feel that the measure- 
ments of the first segment are dependable. Another difficulty 
in getting dependable formulae is the very slight difference in 


Annals Entomological Society of America [Vol. IV, 

the length of some of the segments compared to each other. A 
difference of only two or three microns is all that is found in 
some of the segments. It would be exceedingly difficult to 
eliminate inaccuracies to the extent that the formulae would 
not ba changed by them. Or in other words, the limit of error 
is so small that error, even with the greatest care, is bound to 
occur. Unconsciously the measurements would be made to 
conform to a given formula or to other measurements. The 
writer continually met this difficulty. The measurements 
given in Tables 6-10 were taken with an ocular micrometer at 
a magnification of 660 diameters. Much more accurate work 
was possible than would have been with a camera lucida. 
TABLE 6. PsEUDOcoccus crawii Coq. 


































































• VI 





















































8321 (.57)46 

An examination of the tables mentioned will show the great 
variation met with. Following herewith is a discussion of the 
tables of each species: — 

Crawii Coq. Formula (2, 3, 8) 5, 4, 7, 6, 1, Coq. West 
Am. Scientist '89. 


Specific Characters in Pseudococcus 


There is no question about the determination of this species. 
They were all taken from one plant of white sage {Raniona 
polystachia) at Santa Paula, California. In the tables a formula 
is found which agrees with the one given by Coquillett. In but 
one specimen was the formula the same for the right and left 
antennae of the same individual. All the other formulae 
differed as much as the specifically diagnostic formulae pub- 
lished for all the species of Pseudococcus. 

LoNGispiNus Targ. Formula, (2, 3, 8) (1, 4, 5, 6) 7. New- 
stead, "British Coccidae." Vol. II. The specimens examined 
were taken from palms in the Horticultural Forcing-house of 
Cornell University. Of the ten specimens measured, the for- 
mulae of the right and left antennae of but one specimen were 
identical. No formula was found which agreed with the one 
given by Newstead. The formulae varied as much as the 
specifically diagnostic formulae published for all the species of 

TABLE 7. Pseudococcus 



























































83 (21) (75)64 















































82(31) (75)64 
























Annals Entomological Society of America [Vol. IV, 













































82(13) (56)47 




































































Agrifoliae Essig. No formula is given in the description. 
The description refers however to the figures of the antennae 
for the relative length of the segments. Measuring the figure, 
the following formula is constructed, 7, 1, 3, 2, 4, 6, 5. The 
figure is evidently taken from a nymph, as this species has nor- 
mally eight segmented antennae. These specimens were all 
taken from a single oak tree {Quercus agrifoliae) at Santa 
Paula, California, and are from the lot of specimens that the 
type of the species was taken. In two specimens the for- 
mulae of the right and left antennae were found to be the same 
although the formula of each specimen is different. The for- 
mulae of the twenty antennae varied as much as the specific- 
ally diagnostic formulae published for the species of Pseudo- 


Specific Characters in Pseudococcus 



Pseudococcus citri Risso. 





















81 (23) (67) (45) 











82(13) (67)45 













































































82(13) (45)76 













CiTRi Risso. Formula 6, 3, 2, 1, 5, (4, 6, 7) Newstead, 
"British Coccidae" Vol. II. The specimens of this species 
were taken from coleus in the Horticultural Forcing-houses of 
Cornell University. No formula was found that agreed with 
the one given by Newstead. In one specimen the formulae for 
the right and left antennae were identical. The formulae of 
the twenty antennae varied as much as the specifically diag- 
nostic formulae published for all the species of Pseudococcus. 

Obscurus Essig. Formula 8, 1, 3, 2, 4, 7, 5, 6. Essig, 
"Pomona Jour. Ent." '09. The specimens of this species were 
taken from an elder tree (Sambucus glaiica) at Santa Paula, 
California, and are from the lot of specimens that the type for 
the species was taken. In one specimen the formulae of the 
right and left antennae were identical. 


Annals Entomological Society of America [Vol. IV, 

TABLE 10. PsEUDOcoccus obscurus Essig. 

































■ III 


























































































No formula was found agreeing with the one quoted. In 
specimen X the right antenna has nine segments and in spec- 
imen IX the left antenna has six segments. An examination of 
Table 1 shows that this specimen is the largest specimen studied, 
and as it was found in an egg mass it was undoubtedly an adult. 
Specimen I, which will be seen to be of normal size (Table 1), 
also had the left antenna with six segments. This variation in 
the number of segments was also noted in other specimens. 
The formulae of the twenty antennae measured varied as much 
as the specifically diagnostic formulae for all the species of 
Pseudoc6ccus, as well as one formula placing the specimen in 
the genus Phenacoccus and two formulae placing the specimens 
in the genus Ripersia. 

From the above review it will be readily seen that the rela- 
tive length of the segments of the antennae are valueless as 
specific characters. Other workers as well have found this 

1911] Specific Characters in Pseudococcus 319 

character very variable. Kellogg & Smith, '04, found that in 
twenty-five specimens of Cerepicto yuccae, a closely allied 
genus, no two formulae agreed, "and that there was practically 
as much variety in these formulae as there is among the eleven 
formulae published as specifically diagnostic for eleven North 
American species of the genera Cereputo and Phenacoccus." 
Again Tinsley, J. D., 'OS, in discussing the variation in the 
antennae of P. virgatus Ckll. gives eleven different formulae. 
Again the same author, 1900, in his description of a new species, 
P. texensis Tins., gives three different formulae. Ehrhorn, 
Edw. M., 1900, in his description of a new species, P. maritimiis 
Ehr., gives four formulae. The universal use of antennal for- 
mulae in descriptions is unfortunate, as this has given an excuse 
for the creation of new species and is valueless in analytical 
tables for the determination of specimens. The sooner that 
the valuelessness of this character is realized by systematists 
describing new species of this genus, the sooner will a search for 
valuable characters be begun and a serious mistake in taxonomy 
be corrected. 

Legs. Tables 11-15. The length of the legs is used to 
quite an extent in descriptions. Their length compared to the 
length of the antennae is often stated, also the length of the 
dift'erent segments compared with each other. They are often 
spoken of as long and strong. The presence of hairs is often 
noted, also that of knobbed digitules. 

For the study of the legs five specimens of each species were 
used. They were the same specimens that were used in the 
study of the antennae. The measurements of the different 
segments are the greatest length of these segments, so that the 
sum of the lengths of the different segments will be greater than 
the length of the leg. Like the basal segment of the antennae, 
the coxa is very difficult to measure and the results are not 
entirely reliable. 

At the right hand side of the tables, formulae are 
appended. These formulae are constructed in the same man- 
ner as the antennal formulae. The segments are numbered in 
order beginning with the coxa. 

The formulae show but little if any more satisfactory results 
for specific determination than do the formulae of the antennae. 
The formulae of each pair of legs for each of the species will be 
discussed together. 

320 Annals Entomological Society of America [Vol. IV, 

Prothoracic Legs. In crawii Coq. and citri Risso the third 
segment is always the longest, the fourth segment coming next 
in order. In the remaining three species the third segment is 
usually the longest but may be equal to or less than the fourth 
segment. The first segment is always third in order in the 
formulae. In no species does the second segment bear any fixed 
relation to the fifth. The sixth segment is always much the 
shortest and comes last in order iq^ the formulae. 

Mesothoracic Legs. In crawit Coq. the third segment is 
always the longest. In the other species the position of the 
third and fourth segments vary in relation to each other. The 
first segment always comes third in the formulae. The second 
and fifth segments vary in relation to each other in each of the 
species. The sixth segment always comes last in the formulae. 

Metathoracic Legs. In obscurus Essig and citri Risso the 
fourth segment is usually longer than the third. In obscurus 
Essig an exception is seen to this in the right leg of Specimen 
XL In obscurus Risso an exception is seen to this in specimen 
VII. In the other species the fourth segment is always longer 
than the third. The first segment always comes third in the 
formulae. In no species do the second and third segments bear 
any fixed relation to each other. The sixth segment is always 
much the shortest and comes last in the formulae. 

It will be seen from the above discussion that the variation 
in the formulae is too great for them to be of service in specific 
determination. Exceptions are found to any generalization 
that might be made. The limits within which there can be 
variation are so small that the variations are almost sure to 
go beyond these limits. Other parts of the legs as setae, 
digitules, etc., apparently offer no characters of a specific 


Specific Characters in Pseudococctis 



5 IL 
























































































o ■ 






















































































Annals Entomological Society of America [Vol. IV, 

TABLE 12. 












































































































































































Specific Characters in Pseudococciis 


TABLE 13. PsEUDOcoccus longispinus Targ. 





































































































































































Annals Entomological Society of America [Vol. IV, 

TABLE 14. PsEUDOcoccus obscurus Essig. 



































































































































































Specific Characters in Pseudococcus 


TABLE 15. Pseudococcus agrifoliae Essig. 








































































































































































Marginal Wax Filaments. The number, length and 
stoutness of the marginal wax filaments is a character often 
used. The length of the filaments compared to the length of 
the body and to each other is the form most used. The favor- 
ite character for separating longispinus Targ. and citri Risso 
is the extreme length of the caudal filaments in the former. 

326 Annals Entomological Society of America [Vol. IV, 

This character refers to specimens before treatment with 
caustic potash. Only a slight study has been made of this 
character because living specimens of three of the five species 
studied were not available. A difference however has been 
noted between citri Risso and longispinus Targ., in the 
former the appendages are robust and covered with granules of 
wax, in the latter the appendages are slender and do not 
have granules upon them, being more linear and wire like. 
The objection to the use of this character in the deter- 
mination of species is the ease with which the filaments are 
removed and the danger that the specimens consequently 
would not be in a natural condition. Also in recently moulted 
specimens the normal waxy covering would not be formed. 
The length of the caudal filaments in longispinus Targ. is evi- 
dently a good character if the specimens can be examined in a 
normal condition. 

Anal Lobes. — The anal lobes are frequently mentioned as 
being faintly indicated, normal or prominent. Also the presence 
of one or more setae is occasionally used. 

The observations made upon the segmentation of specimens 
would apply very largely to a discussion of the anal lobes. As 
a specimen becomes older and more distended with eggs the 
anal lobes become less prominent. Consequently in the use of 
this character the age of the individual would have to be care- 
fully considered. In the setae of the anal lobes there is evi- 
dently a good character. (Tables 1-5). Upon each anal lobe 
there are generally several setae. One of these, however, is 
much longer and more robust than the others and is usually situ- 
ated at the caudal extremity of the lobe. This is the seta referred 
to in the tables and discussion. The variation in the length of 
this seta is considerable, but the limits of variation are so great 
that this does not lessen the value of the character. The com- 
parison of the length of this seta to the length of the setae of the 
anal ring is probably the most useful way of expressing the 

Setae of the Anal Ring. — The number of setae on the anal 
ring is a generic character and although often given in descrip- 
tions is of no importance as a specific character. Their length 
compared to some other part of the insect is seldom mentioned. 

However, the length of these setae is a very promising 
specific character. The variation is considerable but not so 

1911] Specific Characters in Pseudococcus 327 

great as to nullify the usefulness of the character. To deter- 
mine the distal termination of the setae is sometimes difficult, 
but it can be done with considerable accuracy. Another 
difficulty is that the distal end of the setae have a tendency to 
curl, but in spite of this and the above mentioned factor the 
length of a seta can usually be ciuite accurately measured. As 
stated above the comparison of the length of the setae of the 
anal ring with the setae of the anal lobes is probably the most 
available way to use this character. 

In agrijoliae Essig considerable variation is found and the 
character is hardly satisfactory. The setae of this species are 
especiall}' difficult to measure because they are not robust. 

Dermis. — The presence of setae and groups of spinnerets 
upon the dermis is a character commonly noted. This is 
usually not given enough in detail to be of assistance in the 
separation of species. 

Ec;(;-Sac. — Considerable use is made of the egg laying 
habits of the species. Whether the egg-sac is of a fibrous 
nature or is a mealy secretion is often stated. Also its presence 
or absence is noted, depending whether the species is ovo- 
viviparous or oviparous. 

If a study of specimens of Pseudococcus can be made with 
the insects in a natural condition, the manner of oviposition is a 
good specific character. Whether a species is ovoviviparous or 
oviparous is not always a sharply separated condition, for an 
individual may give birth to living young and also deposit 
eggs. However, the formation of an egg sac is an indication 
that a species is oviparous and the formation of this egg-sac is 
constant for the species. In a normally ovoviviparous form a 
mealy secretion which is distinctly of a fibrous nature is made 
upon the plant. Crawii Coq. and longispinus Targ. do not 
form a distinct egg-sac but form a mealy secretion upon the 
plant. Citri Risso and obscurus Essig form an egg-sac of con- 
siderable size. Agrijoliae Essig forms an egg-sac but not so 
extensive a one as the last two named species. 

The writer is working on a monograph of the North Amer- 
ican species of this genus. He would be glad to exchange 
specimens of Coccidae for species of Pseudococcus not now in 
his collection. Correspondence to this end is invited. 

328 Annals Entotnological Society of America [Vol. IV, 



By Charles H. T. Townsend, 
Piura, Peru. 

In a paper about to be published re\aewing the results set forth in 
Pantel's 1910 publication in La Cellule, I stated that the said results 
were entirely unknown to me at the time of sending in the last proofs 
and corrections, including the addenda, to my paper in the June (1911) 
issue of the Annals of the Entomological Society of America. 
Largely due to this fact but also in part to a regretable haste to present 
as complete a statement of results to date as possible, I have com- 
mitted several errors that must be corrected. Aside from certain 
typographical errors which will be apparent, the following brief state- 
ments will serve to cover the points in question: 

I should have stated that the spermathecae are "usually" three in 
number. Pantel has shown that Chaetotachina rustica has only one 
spcrmatheca, and Siphona (Pantel et al.) has onl}' two. (Page 127.) 

The uterovagina is a vaginal tube which functions anteriorly as a 
fertilizing but non-incubating uterus when the incubating uterus is 
absent. It may be pointed out that fertilization is a true function of 
the muscoid uterus, quite as much so as incubation. The eggs 
must be held for a certain time to insure fertilization. (Page 127.) 

Phasia has no uterus and deposits flattened macrotype eggs on 
host (Pantel). Alophora has likewise no uterus, but evidently deposits 
its elongate eggs subcutaneously in host (Pantel). Each is the type of 
a distinct tribe or group vmit. (Page 128). 

Compsilura, Eucelatoria and allies have a separate piercer and 
larvipositor in the female. Pantel has first properly described these 
organs in Compsilura, and I have verified his results in Eucelatoria. 
The larvipositor is a short subconical membranous tube approximated 
to the upper base of the elongate curved piercer. The maggots arc 
ejected through the larv'ipositor within the puncture in the skin of the 
host made by the piercer. This is more fully explained in the paper 
above mentioned reviewing Pantel's results, where are also given the 
functions of the ventral carina of the female. (Pages 130, 140.) 

The Peruvian species which I referred to Tricholyga proves to be a 
typical Euphorocera, truly congeneric with E. tachinomoides T. of the 
Southwest. (E. claripennis of Coqt. is not a Euphorocera.) (Page 131.) 

It is probably a mistake to suppose the existence of a special mem- 
branous anal pad for attachment of the maggots of the Hystriciinae to 

1911] Annas Entomological Society of America 329 

plant surfaces. This is evidently effected by the chorion in whole or 
part, probably including the vitelline membrane. (Pages 131, 132, 
133, 134.) 

The Myiophasiine flies are parasites of weevil grubs in buds as well 
as fruits, as for example cotton buds (squares) and doubtless buds of 
other plants. (Page 136.) 

It seems probable that the Cuterebrine flies have a uterus in which 
the eggs are partially incubated. The hinged lid of egg is at the 
cephalic end. (Page 137.) 

The piercer-like organ of the female of Emphanopteryx is probably 
functional in piercing the skin of the host, but this has yet to be 
demonstrated. (Page 140.) 

Celatoria probably has the piercer and lar\npositor as separate 
organs. (Pages 140, 141.) 

Phasiatacta has the chorion with alveolae (not areoles) gathered 
around a dorsal opaque area. (Page 144.) 

Pyrrhosia evidently can not go in the tribe Eumyobiini, since 
Pantel has shown that it has the eggs and maggots regularly arranged 
in the uterus. (Page 147.) 

Certain of the above points are brought out fully in the paper 
referred to as reviewing Pantel's results, besides many other important 
points in connection with the subject. 

A certain number of the series given represents subfamilies, but the 
great majority corresponds to tribes. Each series name given in the 
paper should have the final "E" changed to "I" and serve as the 
name of a tribe, each tribe taking as its type the type species of the 
genus from which its name is derived. At least seven families and 
twenty subfamiHes should be recognized in the Muscoidea on the basis 
of our present knowledge, but it is premature to attempt to define 
these at present, for further knowledge of the many fonns yet to be 
investigated will quite certainly modify our present conceptions. 

The article in "Science" on "Muscoid and especially Tachinid 
Synonymy" appeared in the issue for June 2, 1911. This and the 
forthconmig review of Pantel's 1910 paper, which review also includes, 
correlation of results secured by Portchinski a quarter-century ago,, 
taken with the present corrections and the paper to which they refer,, 
will present a fairly complete general statement of the progress of this, 
work up to the early part of the present year. So far as known to me. 
Further and more extended papers are in preparation. 

July 31, 1911. 

330 Annals Entomological Society of America [Vol. IV 


The next annual meeting of the Entomological Society of America 
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The annual ineeting of the American Association of Economic 
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Hilton, W. A. — The Structure of the Central Nerv- 
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Forbes, W. T. M. — A Structural Study of the Cater- 
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Palmer, Miriam A. — Some Notes on Heredity in the 

Coccinellid Genus Adalia Mulsaut 283 

Smith, P. E. — Specific Characters Used in the Genus 

Pseudococcus 309 

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Volume IV. Number 4. 


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DECEMBER, 191 1 



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The Entomological Society of America 

Volume IV DECEMBER, 1911 Number 4 


By David T. Fullaww. 

All the gall-making species of the hymenopterous family 
Cynipidae are included in the natural group or division 
Cynipinse. Other members of the family, which is well repre- 
sented in California, are parasitic on dipterous, coleopterous, 
and wood-boring hymenopterous larvae.* The gall-making 
species have been collected and studied by a number of American 
students, including Osten-Sacken, Bassett, Ashmead, Gillette 
and others, but previously no thorough systematic collecting 
of the galls or flies has ever been attempted to the writer's 
knowledge, and the descriptions of the California species are 
scattered through the various entomological periodicals of the 
past thirty-five or forty years. 

In 1906, Miss Rose W. Patterson, (now Mrs. C. B. 
Blakeman), a student of entomology in Stanford University, 
under the direction of Professor Kellogg, began a systematic 
collection of the galls occurring in the vicinity of Stanford 
University and of San Jose, California, which extended through 
several years, the range of her collecting being widened on 
several occasions by excursions into the northern part of the 
state. To these collections there were added the contributions 
of students and other interested persons from different sections. 
Specimens bred from this material were carefully labelled and 
preserved by Miss Patterson with voluminous notes, but her 
removal from the university prevented the completion of the 
work of identification and description, and the whole collection, 
was recently turned over to the writer to be worked up. The 

* They are also recorded from Hemerobius and Aphidaj. 


332 Annals Entomological Society of America [Vol. IV, 

Stanford collection, which includes eighteen undescribed spe- 
cies, forms the basis of the present work, in which it is 
attempted to bring together in monographic form all the Cali- 
fornian species. The table for genera and generic definitions 
have been adapted from Dalla Torre and Kieffer's monographs, 
from which much of the nomenclatorial data has also been 
derived. The author is greatly indebted to Professor Kellogg, 
under whose direction the work was performed, and to Mr. 
William A. Beutenmuller and Professor C. P. Gillette, who 
have compared specimens of doubtful identity with types in 
their possession, for helpful suggestions and advice during the 
progress of the work. 

Fam. CYNIPID.4];. 
Subfam. cynipin.^;. 

Cynipidcs, Pscnides, Inquilin;c. T. Hartig, Zeits. f. Ent., vol. 2 (1840), 
p. 187,' 1<)7. 

Cyniphoidca-. A. For.ster, Vcrh. Zool. Ges. Wien, vol. 10 (1860), p. ,320, no. 2. 
Cynii)ina. C. G. Thomson, Opusc. Ent., vol. 8 (1877), p. 778. 
Cynipinje, Inquilinae. Ashmoad, Tr. Am. Ent. Soc, vol. 1,3 (1886), p. 60. 
Cynipina;. Dalla Torre, Cat. Hymen,, vol. 2 (1893), p. 37. 

Body rugose, shagreened or punctate, rarely entirely smooth. 
Scutcllum without cu]3S, sometimes with deep impression on disc but 
not ctip-shaped. Wings usually with three inore or less complete 
cubital cells, cubitus arising in the middle of the basal vein (in one 
species wanting altogether). Areolet closer to base of radial cell than 
middle. Second segirient of posterior tarsi without spine. Four first 
abdominal tergites of unequal size, second nearly always at least half 
as long as abdomen. Stemites ordinarily more or less visible. Hypopy- 
gium usually teminating in a point, ventral valve at least as long as 
broad, sometimes plowshare-shaped. Phytophagous species, living 
in galls. 


1 Wings more or less foreshortened, not reaching beyond the middle of the 

abdomen 2 

Wings normally developed 3 

2 Thorax covered with a dense pubescence, flat, closely punctate; mcsonotum 

glabrous in the middle; antennae 12-segmented. Galls on QKeniis. . . . 

5 Trichoteras .\shm. 
Thorax only sparsely pubescent, evenly rugose or wrinkled; antennae 13-14 
segmented. Galls on Quercus 1 Biorhiza Westw. 

3 1st abdominal segment longitudinally striate, 2d and 3rd segments con- 

nate; face radiately striate; ventral valve short. Inquilines in galls on 

Quercus 9 Synergus Hartig 

1st abdominal segment smooth 4 

4 Face with two parallel ridges from insertion of antennae to clypeus; 

antennae 9 12-14 segmented, d' 14-15 segmented; scutellum with liasal 
fovea?; radial cell closed. Inquilines in galls on Quercus. . 10 Ceroptres Hartig 
Face without such ridges 5 

1911] Cynipidce of California 333 

5 Hypopygium plowshare-sliaped, tarsal claws simple 6 

Hypopygium not usually produced, truncate and ending usually in a short 

spine (ventral valve) 8 

6 Scutellutn without foveae; radial cell open on costal margin; abdomen 

microscopically reticulate. Galls on Rosa 15 Lytorhodites Kieff. 

Scutellum usually with basal foveaj; abdomen without microscopic 
reticulation 7 

7 Radial cell closed. Galls on Rosa 14 Rhodites Hartig 

Radial cell open. Galls on Quercus 13 Compsodryoxenus Ashm. 

8 Suture separating mcsonolum and scutellum wanting, the latter without 

fovejE, anterior margin not elevated in a ridge, an arcii:iie transverse 
groove delimiting mesonotum posteriorly, parapsidal grooves wanting 

or not distinctly percurrent. Galls on Quercus 2 Neuroterus Hartig 

Suture separating mesonotum and scutellum, anterior margin of latter 
elevated to form a ridge & 

9 Body covered with silky pile, aVjdomcn dorsally sometimes glabrous; 

radial cell open. Galls on Quercus 6 Cynips L. 

Abdomen glabrous, 2nd segment alone sometimes sparsely pubescent 
laterally at base 10 

10 Radial cell closed; pronotum not medially contracted 11 

Radial cell open or partly open on costal margin; pronotum sometimes 

medially contracted 12 

11 Mesonotum wholly smooth. Galls on Rubus 12 Diastrophus Hartig 

Mesonotum not wholly smooth; antennae 12-segmented. Intjuilincs in galls 

on Rosa and Quercus 11 Periclistus Forst. 

12 2nd abdominal segment produced linguiform on dorsum; parapsidal grooves 

percurrent; base of scutellum with an arcuate transverse groove; ridges 

on metanotum arcuate; claws bidentate. Galls on Quercus 

3 Diplolepis L. Geoffr. 
2nd abdominal segment not produced linguiform 13 

13 Claws simple, sometimes ob.scurely dentate, but then antennae slenderer 

at apical third than at middle, and metanotal ridges angularly curved. 

Galls on Quercus 8 Callirhytis Forst. 

Claws bidentate; antennae not slenderer at apical third than at middle; 
metanotal ridges straight and parallel, or arcuate 14 

14 Scutellum l:)asally with arcuate transverse groove; parapsidal grooves 

incomplete; head and thorax densely pubescent; Galls on Quercus 

4 Disholcaspis D. T. & Kieff. 
Scutellum with basal foveae; parapsidal grooves percurrent. Galls on 
Quercus' 7 Andricus Hartig 

1 BIORHIZA Wcstw. 

Biorhiza, Westwood, Intr. Classif. Ins., vol. 2 Syn. (1840), p. 56. 
Philonix, A, Fitch, 5th Rep. Ins. N. York (1859), p. 3. 

Wings in agamic generation wanting or very rudimentary, in sexual 
female very rudimentary, in male always present. Antennae of female 
13-14 segmented, of male 15-segmented, 3rd segment longer than the 
4th, in male often strongly excised, succeeding segments progressively 
shorter but all longer than broad. Pronotum narrow in the middle, 
mesonotum in wingless generation only partly smooth and shining, in 
generation with developed and rudimentary wings usually entirely so, 
parapsidal grooves complete or little marked, scutellum in wingless 
generation with a transverse groove at base, in female with rudimentary 
wings, with weak foveas separated by a carina, in generation with devel- 
oped wings, with two sharply separated foveae. Abdomen large, lat- 
erally compressed or globose, smooth or pubescent, ventral spine short. 
Tarsal claws bidentate. 

334 Annals Entomological Society of America [Vol. IV, 

Biorhiza califomica (Beutenm.) 
Philonix califomica, Beutenmiiller, Ent. News, vol. 22 (1911), p. 69. 
"Female. Head pitchj^ brown black, minutely rugose with scat- 
tered, short hairs. Antenna; 13-jointed; first joint stout, cj'lindrical ; 
second joint shorter, stout and rounded at the tip ; third joint very long 
and slender; fourth, fifth and sixth joints slender and shorter than the 
third; remaining joints gradually becoming shorter and thicker toward 
the thirteenth, all pitchy brown and pubescent. Thorax pitchy brown 
or dull rufous, evenh^ rugose, somewhat wrinkled and with a few scat- 
tered hairs. Parapsidal grooves very fine and somewhat lost in the 
rough surface anteriorly, convergent at the scutellum. Scutelltun 
evenly rugose like the thorax, and of the same color. Abdomen com- 
pressed, convex at the sides and rather sharply keeled on the dorsum 
and venter, dark pitchy brown, smooth and shining. Legs pitchy 
bro's^Ti, somewhat paler than the abdomen and pubescent. Wings 
aborted, not extending to the middle of the abdomen. Length 1 mm. 
^ "Gall. On the upper surface of the leaves of a species of white oak. 
Monothalamous. Rounded, flattened disc-like, becoming slightly 
elevated toward the middle. The sides are flat and very thin, and the 
gall rests closely on the leaf. The larva lives in the center of the elevated 
part. The color is pinkish or purplish, with the apex sometimes j^ellow- 
ish. Width, 3 to 4 mm. Height, 1 mm. 

"Habitat. Kern Co., California, January." 

(Wm. Beutenmiiller.) 

I have not seen specimens; the type is in the National 


C>Tiips (part.), Linne, Syst. Nat. ed. 10 (1758). p. 343, 553. 

Ncuroterus, Spathegaster, Hartig, Zeits. f. Ent., vol. 2 (1840), p. 1S5, 192, 
186, 194. 

Ameristus, A. Forster, Verb. Zool. Ges. Wien, vol. 19 Abh. (1869), p. 330, 3.33. 

Dolichostrophus, Ashmead, Tr. Am. Ent. Soc, vol. 14 (1887), p. 129 nota. 

Head, thorax and scuteUum smooth or microscopically retictdate 
and shining. Parapsidal grooves wanting or very indistinct. Meso- 
notum not separated from scutellum by a suttire, posterior margin with 
an arcuate recess or indention, a broad transverse groove at base of scu- 
tellum. AntenncB of female 13-14 segmented, of male 14-1.5 segmented. 
Abdomen large and subpetiolate in female, small and with a long petiole 
in male. Wings of male verj- long, usually shorter in female. Radial 
cell nearly alwaj^s open and very long. In galls on Quercus. 

Neuroterus quercus-batatus (Fitch). 
Cynips quercus-batatus, A. Fitch, 5th Rep. Ins. X. York (1859), p. 30. 
Neuroterus batatus, G. Mayr, Gen. Gallenb. Cynip. (1881), p. 37. 
Neuroterus quercus-batatus, Dalla Torre & Kieffer, Das Ticrreich, lief. 24 
Cynipida (1910), p. 334. 

Female. Black, shining, mouth-parts, base of antenna?, legs 
bcA'Ond tibiae and at joints above, tegiJa?, pedicle and ovipositor brown- 
ish. Head faintly rugose, face pubescent, antennae 13-segmented, 

1911] CynipidcE of CalifoDiia 335 

fuscous to black, except three first segments, which are brown, only 
slightly pubescent, 1st and 2nd seginents stout, 2nd the same width 
throughout, 3rd segment longest, not as 1 and 2 together, 3rd and 4th 
very slender, becoming thicker and shorter outwardly to 7th, S-12 
subequal, last only a little longer than penultimate. Thorax micro- 
scopically reticulate, mcsonotum withoixt parapsidal grooves but with two 
rather large basal depressions separated by a median ridge, scutellum 
without basal foveje, smooth, sparsely pubescent, sculpturing on 
mesonotum and scutellum excessively fine, on pleura and prothorax 
somewhat coarse. Abdomen smooth, shining, much compressed, about 
as broad as long, ovipositor exserted. Wings hyaline, pubescent, 
subcostal, radial, basal and cross-veins distinct, black, radial cell long, 
open, vein at base angulate, areolet distinct but small, cubitus indis- 
tinctly reaching basal. Length 1.75 mm. 

iNIale. Microscopically reticulate or rugose, rather shiny. Head 
black, ocelli, mouth and antenna; at base brown, the latter 14-segmented, 
fuscous to black from 4th seginent outwardly, 3rd a little longer than 
1 and 2 together and excised at distal end, 4th about two-thirds of 3rd 
and subequal with oth, following segments a little smaller and subequal 
except 13th, all somewhat pubescent. Prosternum blackish, pronotum 
narrow in the middle, at sides yellowish brown, reticulate and shining, 
the color extending up on to the scapula?, mesothorax and scutellum 
smooth, shining, microscopically sculptured, the latter fuscous yellowish 
brown, mcsopleura fuscous brown, coarsely sculptured, shining, meta- 
thorax sordid white, slightly rugose. Abdomen long petiolate, smooth, 
shining, at base sordid white to yellowish white, otherwise black, much 
compressed apicaUy and pubescent at tip. Legs very pale yellowish, 
tips of tarsi black. Wings extending be^-'ond abdomen more than its 
length, hyaline, pubescent, radial cell long and o]3en, vein at base arcuate, 
cubital vein reaching basal, areolet distinct and large. Length 2 mm. 

Gall. Early summer galls on under side of leaf of Quercus douglasi, 
a flat, irregular swelling, distorting the leaf, polythalamous, S mm. by 
5 mm., and a sordid brown color. Late summer galls, from which adult 
flies emerge the following spring, small hard woody swelling in tenninal 
twigs of Q. lohata, containing numerous long, oval larval cells imbedded 
in soft spongy interior of gall. 

Habitat. Stevens Creek, be^^ond Cupertino, Cal. (R. W. 
Patterson.) San Jose, Cal. (Rose Patterson.) 

Neuroterus saltatorius (Riley). 

Cvnips saltatorius (Hy. Edwards in MS), C. V. Rilov, Tr. Ac. St. Louis, 
vol. .3 (1876), p. 21.3. 

Neuroterus saltatorius, Ashmead, Tr. Am. Ent. Soc, vol. 14 (1887), p. 128. 

Female. Black, smooth and polished, legs at joints sordid white. 
Head microscopically reticulate, antennae 13-segmented, filiform, 
slightly incrassate towards tip, 1st and 2nd segments stout, 3rd longest, 
about twice as long as 2 or 4 but not as long as 1 and 2 together, succeed- 
ing segments subequal, the three last a trifle longer than preceding ones. 
Thorax smooth and flat, faintly sculptured, with two large depressions 

336 Annals Entomological Society of America [Vol. IV, 

posteriorly on either side of median line, which is ridged, scutellum 
rugose. Abdomen as broad as long, much compressed, ovipositor 
exserted. Wings hyaline, pubescent, veins blackish, radial cell long, open, 
vein at base slightly angulate, areolet present but two of the enclosing 
veins indistinct, cubitus not reaching basal vein. Length 1.5 mm. 

Gall. Small, subglobidar, 1.25-1.50 mm., pale reddish brown, with 
a small nipple at either pole; slightly sculptiu-ed. 

Habitat. Stockton, Cal. (Hughes Ranch.) 

3 DIPLOLEPIS L. Geoffr. 

Cvnips (part.), Linne, Syst. Xat. ed. 10 (175S), p. 343, 553. 
Diplolepis (part.), L. Geoffrey, Hist. Ins., vol. 2 (1762), p. .309. 
Drvophanta, A. Forster, Verb. Zool. Ges. Wien, vol. 19 Abh. (1869), p. 331, 
334, 33o. 

Cheeks not more than half the length of the eye. Parapsidal grooves 
pcrcurrent. ScuteUtun without fove«, an arcuate transverse groove 
at base, rarely interrupted in the middle. Metanotal ridges curved. 
Radial cell open at the margin. Tarsal claws usually bidentatc. Abdo- 
men longer than ^•ertically broad, 2nd tergite produced caudally lingui- 
form. Head and thorax in agamous generation densely pubescent, 
antennas 13-segmented and clothed with long, erect hairs. Body in 
sexual generation for the most part glabrous, antenna; of female 14-seg- 
mented, without long erect hairs, of male 15-segmented. Mesonotum 
smooth and shining. Abdomen in male petiolate. In galls on Quercus. 

Diplolepis discus (Bass.) 
Drvophanta discus, H. F. Bassett. Tr. Am. Ent. See, vol. 26 (1900), p. 326.' 
Diplolepis discus. Dalla Torre and Kieffer, Das Tierreich, lief. 24 Cvnipidae 
(1910), p. 362. 

"Head black. Antcnnte thirteen jointed, joints one and two rather 
large, subcqual, third long, fourth two-thirds as long as the third, 
remainder gradually shorter, all j^ellowish red. Thorax smooth, 
shining, with a few scattered hairs and deep parapsidal grooves. Scutel- 
lum slightly rugose. Foveas not distinct. Abdomen dark, shining 
brown. Legs dark brown. Wings rather large; veins very pale, almost 
colorless. Areolet wanting. Cubitus nearly obsolete. Radial area 
open. Body .06, antenna .05, wings .07. 

"Galls. Among the galls sent me several years ago by Mrs. E. H. 
King, from Napa City, California, were a few specimens from which no 
insects appeared, but from which I removed three dead but perfectly 
developed individuals. The galls were circular, flat, sessile discs growing 
in clusters on the under sides of the leaves of some species of oak, closely 
resembling Q. alba; but I am not sure this oak grows in that section. 
The galls are hardly one-eighth of an inch in diameter, and except in 
size and color might be taken for what is, I think, called the 'blue 
spangle gall,' not uncommon on the white oak in the Atlantic States. 
It is smaller and lacks the blue color." (H. F. Bassett). 

I have not seen examples of this species. 

1911] CynipidcE of California 33/ 

Diplolepis clavula (Beutenm.) 
Dryopianta clavula, BeutcnmuUer, Ent. News, vol. 22 (1911), p. 67. 
Female. Reddish brown, eyes, ocelli, mandibles, oral margin, 
occiput, a broad median stripe from occipital margin to the mouth 
(broken beneath ocelli), antenna2, prostemum, pronotum abo^'e and 
below, dorsal and subdorsal vitt» on mesonotum concurrent with 
median longitudinal lines and smooth lines over base of wings, meso- 
pleura, base of scutellum and a median spot, metanotum and abdomen 
wholl}^ black. Head reticulately rugose, face pubescent, antenna 
14-segmented, 1st segment stout, clavate, 2nd oval, 3rd long but not 
as long as 1 and 2 together, 4th as long as 3, succeeding segments to 9th 
progressively shorter, 9th and following segments subequal except the 
last, which is longer than penultimate by one-half, all rather pubescent. 
Pronotum narrow in the middle, rugoso-punetate, pubescent, mesono- 
tum faintly rugose and deeplj- punctate, each puncture with a pale 
brown hair, parapsidal grooves distinct, reaching anterior margin, 
median longitudinal lines extending half-way to posterior margin, 
smooth lines over base of wings rather short, mesopleura smooth, 
shining black, densely pubescent, a triangular area beneath wings 
aciculate, scutellum rugose, pubsecent, fovese indistinct, a transverse 
arcuate groove at base with median carina, metanotum somewhat 
punctate, pubescent. Abdomen smooth, shining, somewhat compressed, 
the second tergite produced caudally acutely to a point, the posterior 
margin oblique, pubescent at base, 3rd tergite fairly wide, others con- 
cealed beneath, ovipositor sheath expanded at tip, with an apical tuft of 
hairs, ovipositor exserted. Legs fuscous brown and pubescent. Wings 
hyaline, quite pubescent, veins distinct, radial cell long, open at the 
margin, vein at base angulate and clouded with brown, areolet moderate, 
cubitus nearly reaching basal vein, radius inerassate at the tip, cubital 
cell with a large brown cloud at base and numerous spots at apex, a 
brownish cloud beneath 2nd cross vein, one at break in anal cell, and 
another at base of cubitus. Length 2 mm. 

Gall. Small gourd or trumpet-shaped galls on the leaves of Quercus 
lobata, oval or subglobular outwardly, with long neck which is somewhat 
expanded at point of attachment. The gall is 7 mm. long, 2 mm. wide, 
the neck narrowing to less than 1 mm. ; the walls quite thin, the large 
oval larval chamber Ij-nng directly beneath. It has a reddish appear- 
ance and is faintly rugose from minute crystalline bodies lying on the 
surface. It is also often covered with a grayish brown tomentum. 

Habitat. Palo Alto, Cal. (Miss Bertha Wiltz.) Napa and 
Sonoma Counties, California (Beutenmuller). 

Diplolepis echina (O.-S.) 

Cynips echinus, Osten-Sacken, Tr. Am. Ent. Soc, vol. 3 (1870), p. 56. 
Dryophanta echina, Beutenmuller, in litt. 
Dryophanta speciosa, Beutenmuller, in litt. 

Female. Reddish brown, eyes, ocelli, tips of mandibles, oral 
margin, antennae distally from 2nd segment, dorsal and lateral vittae 
on mesonotum concurrent with median pair of lines and lines over base 
of wings, and abdomen dorsally black or blackish. Head faintly rugose, 

338 Annals Entomological Society of America [Vol. IV, 

antennas 14-segmented, filiform, outer third slightly thicker, 1st segment 
stout, obconic, 2nd subglobular, 3rd longest, nearly as long as 1 and 2 
together, 4th and suececding segments to 9th progressively shorter, 
9-14 subequal, less than one-half as long as 3rd, all sparsely covered with 
appressed gra^'ish hairs. Thorax faintly rugose and sparsely, in some 
parts rather thickly covered with appressed yellowish gray pubescence, 
pronotum narrow in the middle, parapsidal grooves on mesonotum dis- 
tinctly percurrent, median longitudinal lines reaching half-way to post- 
erior margin, smooth lines o\-er laase of wings distinct and long, scutcllum 
without distinct fovea?, a rather narrow, arcuate transverse groove with 
smooth shining bottom at base. Abdomen smooth, shining, second 
tergite produced eaudally linguiform almost to apex, pubescent at base, 
dorsal valve and sheath of ovipositor prominent, the latter with apical 
tuft of }'ellowish brown hair. Legs rather stout and clothed with a 
grayish pubescence. Wifigs hyaline, pubescent, veins brownish, 
distinct, radial cell open at the margin, vein at base only .slightly bent, 
radius incrassate at tip just before costal margin, areolct distinct, 
cubitus nearly or quite reaching basal vein, a small brownish cloud near 
base of cubital cell, another beneath areolet, and still another at the 
break in anal vein. Length 2.5-3 mm. 

Gall. Moderately large, reddish, echinus-shaped galls attached to 
the leaves of Qiiercus douglasi. Numerous pointed processes project 
from the more or less globular body of the gall, giving it the character- 
istic echinus appearance. The gall is composed of a crystalline sub- 
stance said to be hardened gallic acid. Monothalamous. About 
12 mm. in diameter. 

Habitat. St. Helena, Cal. (Miss Julia Begley). Placer 
County, California (Osten-Sacken). 

Diplolepis douglasi (Ashm.) 

Holcaspis douglasii, Ashmead, Proc. U. S. Nat. Mus., vol. 19 (1896), p. 127. 

Holcaspis douglasi, Dalla Torre & Kieffer, Gen. Ins. Hvracn. Fam. Cynip. 
(1902), p. 53. 

Dryophanta douglasi, G. Mayr, Verh. Zool. Ges. Wien, vol. 52 (1902). p. 290. 

Diplolepis douglasi, Dalla Torre and Kieffer, Das Tierreich, lief. 2-i Cyn- 
ipidjE (1910), p. 369. 

Female. Very similar to D. echina, from which it can scarcely be 
separated except on the character of the gall. In specimens before me 
I notice the following minor differences : antennas black distally from 6th 
segment instead of froin 2nd; abdomen black on dorsum only posteriorly; 
areolet rather indistinct and only the cloud near base of cubital cell 

Gall. Pink, star-shaped galls occuning on leaves of Qucrcus lobata, 
composed of a crystalline substance similar to the material fomiing the 
gall of D. echina, and covered with a pale bloom which imparts a lilac 
shade to the whole body. The gall is 8 mm. high and 10 mm. in diam- 
eter, the pedestal widening rapidly to the dorsal rim, which bears about 
eight irregular pointed projections fonning the star. Mononthalamus. 
Hollow within, the oval larval chamber partly attached. 

Habitat. San Jose, Cal. (R. W. Patterson.) Marin County, 
California (Beutenmuller) . 

1911] Cynipida of California 339 

Diplolepis dubiosa n. sp. 

Female. Black, the antennte, legs, tips of mandibles, tegulse, post- 
scutellum and sheath of ovipositor luteous. Head faintly rugose and 
covered with closely appressed whitish hairs, antenna; 14-segmented, 
filiform, outer third slightly incrassatc, 1st segment stout, obconic, 
2nd suboval, 3rd longest, 4th and succeeding segments to 9th progres- 
sively shorter, 9th and following segments subequal, except the last, 
which is somewhat longer then ]:icnultimate. Pronotum narrow 
medialh', mesonotum faintly reticulate, shining, parapsidal grooves 
distinct, reaching anterior margin, scutellum deeply rugose, foveas 
distinct, large and broad, contiguous, separated only by a carina. 
Abdomen about as large as head and thorax together, smooth, shining, 
2nd tergite produced caudally almost to apex, slightly pubescent 
laterally near the middle, spine of ventral valve moderate, pubescent. 
Wings hyaline, pubescent, veins pale brown to black, radial cell long, 
narrow, open at the margin, vein at base only slightly bent, areolet 
distinct but the enclosing nervures (except distal one) very delicate. 
Claws unidentate. Length 2 mm. 

Male. Black, the antenna, legs (except coxe), tips of mandibles, 
palpi, tegulas and petiole luteous. Head faintly rugose, face and cheeks 
covered with long, whitish hairs. Pronotum, mesonotum and pleura 
faintly reticulate, shining, metanotum faintly aciculate, pubescent, 
scutellum deeply rugose. Abdomen long, slender, compressed, pube- 
scent laterally on 2nd segment. Antcnnfe 15-segmented, 1st segment 
obconic, 2nd subglobose, 3rd longest, succeeding segments to 13th 
progressively shorter, 13th and 14th stibcqual, 15th short and pointed. 
Length 1.7.5 mm. 

Gall (PI. XXIII, fig. 1). A small, brown, thin-shelled gall arising, 
several together, in the staminate ament of Quercus agrijolia. The gall 
is about 3 or 4 mms. long, rather angulatc and more or less clavate. 

Habitat. Palo Alto, Cal. (R. W. Patterson.) 

4 DISHOLCASPIS D. T. & Kieff. 

Holcaspis, G. Mayr, Gen. d. Cynip. (18S1), p. 9, 35. 

Disholcaspis (n. n. for Holcaspis, preoccupied), Dalla Torre and Kieffer, Das 
Tierreich, lief. 24 Cynipida; (1910), p. 371. 

Agamic generation. Head and thorax covered with a rather dense 
pubescence, abdomen bare, the sides basally slightly pubescent. Cheeks 
less than half the length of the eyes. Antennae 13-15 seginented, 
pubescent. Pronotum narrow in the middle. Parapsidal grooves 
incomplete, not reaching anterior margin. Base of scutellum with a 
transverse groove. Ridges of the metanotum arcuate. Radial cell 
long and open at the margin. Tarsal claws bidentate. Second tergite 
of abdomen not produced linguifonn caudally. Sexual generation 
unknown. Galls on Quercus. 

340 Annals Entomological Society of America [Vol. IV, 

Disholcaspis truckeensis (Ashm.) 

Holcaspis truckeensis, Ashmead, Proc. U. S. Nat. Mus., vol. 19 (1896), p. 127. 
Disholcaspis truckeensis, Dalla Torre and Kieffer, Das Tierreich, lief. 24 
Cynipida; (1910), p. 380. 

"Gall. An irregtdar, inflated, hard, woody gall, over an inch long 
and about half an inch in diameter, issuing from a slit in a temiinal twig 
of Quercus chrysolepis var. vaccinijolia; polythalamous. 

"Gall-fly. Female. Length, 3.4 mm. This species, in color and 
size, closely resembles H. Jicigera, Ashmead, but differs as follows: 
The 14-jointed antennas, except the first two joints, pleura, and meta- 
thorax blackish, shining ; the rest of the insect — except the dorsum of the 
second abdominal segment, which is obfuscated — brownish yellow; the 
head and thorax punctate, and covered with a glittering white pube- 
scence. Abdomen highly polished, bare, except the sides of second 
segment basally; spine of ventral valve short, stout, hairj'. Wings 
hyaline, veins brown, areolet distinct, cubital cell open at base, while 
the basal vein of radial cell is only obtusely angular. 

"Type No. 3080, U. S. N. M. 

"Two specimens, reared December 6th, 1880, from galls 
collected by Prof. J. H. Comstock, in California, October 16, 
1880." (W. H. Ashmead.) 
• I have not seen examples of this species. 

Disholcaspis eldoradensis (Beutemn.) 

Holcaspis eldoradensis, BeutenmuUer, Bui. Am. Mus. Nat. Hist., vol, 26 
(1909), p. 38. 

Female. Testaceous, eyes, ocelli, occiput, anterior margin of face, 
antennae distally from 8th segment, pecten, dorsal and subdorsal vittas 
on mesonotum concurrent with median longitudinal lines and lines over 
base of wings, median vitta on metanotum and dorsal valve black, 
vertex and front of head, tibia; and tarsi of legs, and abdomen dorsally 
fuscous. Head faintly nigose and co\'ercd with dense grayish pube- 
scence, antennte 14-segmented, 1st and 2nd segments stout, 2nd a trifle 
longer than broad, 3rd segment longest, longer than 1 and 2 together, 
4th segment as long as 3rd, succeeding segments to 10th progressively 
shorter, 10th and following segments subequal except the last, which is 
somewhat longer than penultimate ; all the segments sparsely pubescent. 
Thorax coarsely punctate and covered with pubescence, parapsidal 
grooves on mesonotum indistinct, median longitudinal lines reaching 
half-way to posterior margin, smooth lines over base of wings distinct, 
scutellum rugose, foveas inconspicuous, long, narrow and oblique. 
Abdomen smooth and shining, much compressed, 2nd segment reaching 
half-way to apex, pubescent at base, 3rd, 4th and .5th segments fairly 
broad, dorsal and ventral valves thickly pubescent. Wings hyaline, 
pubescent, veins heavy, black ,radial cell long, open at the margin, vein 
at base of cell decidedly angulate, radius itself almost straight, areolet 
rather large, cubitus not quite reaching basal nerviu-e. Claws simple 
Length 2.5-3 mm. 

1911] CynipidcE of California 341 

Gall. Small, brown, cushion-shaped galls, sessile on twigs of 
Qiiercus kelloggi and Qiiercus lohata, hard and woody with the base 
broadly inserted in a slit in the bark. The top is flat and much pitted. 
Length about 4 mm., breadth about 3 mm., height about omm. 

Habitat. Stanford University, Cal. (R. W. Patterson). 
Sonoma County, California. (Beutenmuller). 

Disholcaspis chrysolepidis (Beutenm.) 
Holcaspis chrysolejiiilis. Beutennuilkr, Ent. News, vol. 22 (1911), p. 68. 
Female. Very similar to D. eldoradensis, from which it can scarcely 
be separated, except on the character of the gall. Specimens before me 
show the following differences: parapsidal grooves deep and distinct, 
reaching half-way to anterior margin, pubescence on the thorax 
less abundant. 

Gall. Galls sessile on the twigs of Qucrcus chrysolepis and Qiiercus 
dumosa and massed together around the stem, 15 to 30 in a mass; very 
irregular but more or less cushion-shaped outwardly and rugose, 3-4mni. 
in diameter, this portion of the gall surmounting smooth, lobular 
enlargements, sometimes arranged in a rosette. Sometimes the galls 
are elevated or arranged palisade-like, the outer face rough, the sides 
smooth, and with a median constriction. They are described as red- 
dish or beef colored when fresh. Monothalamous, with a large internal 
pupal cell. 

Habitat. Alma Soda Springs, Cal. (W. R. DudlejO- Placer 
County, California. (Beutenmuller.) 


Trichoteras, Ashmead, Psyche, vol. 8 (1897), p. 67. 
Trichoteras, Ashmead, Psyche, vol. 10 (1903), p. 150. 

Head and thorax closely punctate, opaque, and densely pubescent. 
Antenna? of female 12-segmented, 3rd segment a little shorter and thicker 
than 4th equal to .5th, succeeding segments to 9th becoming gradually 
shorter, 9th to 11th only a little longer than thick, 12th as long as 10 
and 11 together. Disc of the mesopleura bare, smooth and shining. 
Scutellum cushion-shaped, a little longer than wide, with two smooth, 
lunate fovea? at base. Tarsus of hind legs not longer than tibia, claws 
with a tooth at base beneath. 

Trichoteras coquilletti Ashm. 

Trichoteras cociuilletti, Ashmead, Psyche, vol. 8 (1897), p. 67. 

"Galls. Small, brown, sub-opaque, globular galls, averaging from 
6 to 8 mm. in diameter, and internally with a central kernel or larval 
cell held in place by radiating filaments. 

"These galls were collected by Mr. D. W. Coquillett, at Los Angeles, 
California, from the upper surface of the leaves of an unknown oak, 
who forwarded them to the Department of Agriculture, where three 
specimens of the gall-wasp were reared. Structurally and in general 
appearance the galls very closely resemble Dryophanta poHla Bass., but 
the sub-apterous wasp is quite different from that species. 

342 Annals Entomological Society of America [Vol. IV, 

"Agamous female. Length 2.5 mm. Head and thorax ferruginous, 
closely punetate, and very hairy; prostcrnum and pleura blackish; legs 
fusco-piceous, the articulations paler. 

"Antennas 12-jointed, shorter than the body, the scape fully as long 
as the first joint of flagellum, obconical, and inuch stouter, pedicel one 
and one-half times as long as thick, 2nd joint of flagellum distinctly 
longer than either the 1st or 3rd joint; 4th joint of flagellum a little 
shorter than the 3rd, the 5th and following joints gradually shortening, 
the penultimate joint being scarcely longer than thick, the last joint 
fully as long as the first joint of flagellum, or twice as long as the peiiulti- 
mate. Wings abbreviated, narrowed and not extending beyond tip of 
abdomen, the veins dark brown, the marginal cell open, the areolet 
indicated by the union of the surrounding nervures. Abdomen black, 
polished, pubescent along the sides towards base, and as long as the head 
and thorax together, compressed and viewed from the side it is as 
broad as long, the hypopygium armed at tip with a long spine.- 

"Hab. Los Angeles, California. 

"Type No. 3498, U. S. N. M. 

"Described from 3 female specimens bred Nov. 2G and 29 and 
Dec. G, 1S92." (W. H. Ashmead). 

I have not seen examples of this species. 


Cynips (part.), Linne, Syst. Nat. ed. 10 (1758), p. 343, 553. 

Cynips (part.), T. Hartig, Zeits, f. Ent., vol. 2 (1840), p. 185, 187. 

Cynips, Forstcr, Verh. Zool. Gcs. Wien, vol. 19 Abh. (1869), p 331, 335. 

Cynips, G. Mavr, Gen. d. Cvnip, (1881), p. 28. 

Diplolepis (part.), L. Geoflfroy, Hist. Ins., vol. 2 (1762), p. 308. 

Agamic generation. Body densely pubescent throughout, abdomen 
dorsally sometimes more or less bare. Antennae filifonn, 12-15 seg- 
mented, without long, erect hairs. Pronotum narrow in the middle. 
Parapsidal grooves percurrent. Scutellum with two basal foveae, 
separated by a median carina. Metanotal ridges parallel. Radial cell 
open at the margin. Tarsal claws bidentate. 

Cynips canescens (Bass.) 

Holcaspis canescens, H. F. Bassott, Tr. Am. Ent. Soc, vol. 17 (1890), p. 66. 
Disholcaspis canescens, Dalla Torre and Kieffer, Das Tierreich, lief. 24 
Cjmipidae (1910), p. 378. 

Female. Reddish browm, eyes, ocelli, tips of mandibles, antennae 
distally from Sth segment and a spot on 1st, prostemum, 1st abdominal 
segment, tarsi and ventral valve fuscous to black, clothed with pale 
grajash pubescence throughout except abdomen dorsally, which is 
bare. Head faintly rugose, antennas 14-segmented, filiform, 1st and 
2nd segments stout, 3rd segment longest, longer than 1 and 2 together, 
and a little longer than 4th, succeeding segments progressively shorter, 
the last' in ^me specimens indistinctly di\-ided and longer than 
penultimate. Pronotum narrow in the middle, mesonotum ptmctate, 
each puncture bearing a hair, parapsidal grooves reaching about half- 
way to anterior margin, median longitudinal lines extending about 

1911] Cynipida of California 343 

half-way to posterior margin, smooth Hnes over base of wings distinct 
and broad, mesopleura and scutellum punctate, basal foveas on latter 
obsolete. Abdomen compressed, dorsally bare, smooth and shining, 
ridged apically, pubescent at sides and beneath, 2nd segment occupying 
about one-half its length, 3rd tergite distinct, only stemites of following 
segments visible, dorsal and ventral valves prominent and pubescent. 
Wings hyaline, pubescent, veins prominent, radial cell open at margin, 
vein at base angulate, areolet large, cubitus not reaching basal vein. 
Length, 4 mm. 

Gall. Brownish, globular galls, about 10 mm. in diameter, occur- 
ring singly or in clusters on the twigs of Quercus douglasi. Sometimes a 
little irregular in shape, not much roughened, and covered with a 
yellowish brown fuzz. Monothalamous, with a large, globular larval 
chamber, quite distinct from the cortical layer and held in place by 
loose, spongy tisstie. 

Habitat. Hornitos, Cal. (Miss Hazel Engebretsen). 

Cynips corallina (Bass.) 

Holcaspis corallinus, H. F. Bassett, Tr. Am. Ent. Soc. ,vol. 17 (1S90), p. 66. 
Holcaspis corallina, Dalla Torre, Cat. Hymen, vol. 2 (1893), p. 55. 
Disholcaspis corallina, Dalla Torre and Kieffer, Das Tierreich, lief. 24 
CynipidEE (1910), p. 377. 

Female. Very similar to C. canescens, but the pubescence is regularly 
more extensive on abdomen, only a small rhomboidal area dorsally on 
second segment remaining bare. The antennas are brownish through- 
out, the basal segments darker, but light at the joints. The abdomen 
is not compressed apically but rotund, and all the tergites are visible. 

Gall. Pale yello\\4sh, globular galls, about 12 mm. in diameter, 
found on the twigs of Quercus douglasi. The galls are pointed at the 
poles and the surface is roughened by irregular ridges, or bears short, 
blunt tubercles giving it the appearance of coral. 

Habitat. Mt. Diabalo, California. (Harold Morrison.) 

Cynips multipunctata (Bcutenm.) . 

Dryophanta multipunctata, Beutcnmiiller, Ent. News, vol. 22 (1911), p. 67. 

Female. Reddish brown, eyes, ocelli, tips of mandibles, face medi- 
ally, pecten, dorsal and subdorsal vittas on mesonotum concurrent with 
median longitudinal lines and lines over base of wings, abdomen dor- 
sally, sometimes entirely black or blackish. Head broad, bulged 
beyond the narrow eyes, faintly rugose and punctate, with a thick 
covering of pale yellowish pubescence, antennas 14-segmented, 1st and 
2nd .segments brown, following segments brownish black, 1st to 4th 
segments light brown distally, 1st and 2nd segments as usual stout, 
3rd segment longest, longer than 1 and 2 together and a trifle longer 
than 4th, the following segments progressively shorter except the last, 
which is one-third longer than penultimate. Thorax faintly' rugose 
and punctate, pubescent, except on median longitudinal lines which 
are bare, parapsidal groqves complete, median longitudinal lines reach- 
ing more than half-way to posterior margin, smooth lines over base of 

344 Annals Entomological Society of America [Vol. IV, 

wings distinct and rather long, pleura pubescent, mesopleura spotted 
with black, scutellum rugose and punctate, each puncture bearing a 
hair as on head and mesothorax, black at base, fove^ indistinct. 
Abdomen smooth, shining, faintly punctate, all the segments clothed 
with long, straight, pale yellowish pile, but dorsally and laterally bare 
in spots, rotund but ridged dorsally, 2nd segment occupying half its 
length, 3rd and 4th segments also wide, ventral valve large, triangular, 
with broad apex, bearing a large tuft of hairs, dorsal valve also hairy at 
tip. Legs pale brown and clothed with hairs. Wings large, hyaline, 
pubescent, veins brownish, black, subcostal, radial, anal and cross 
veins thickened, the radial vein incrassate at tip, radial cell open and 
rather short, vein at base angulate and clouded, areolet large and dis- 
tinct, cubital vein reaching almost to basal, clouded area at base of 
cubital and discoidal cells and at break in anal vein, the cubital cell 
with 12-1.5 black spots. Length 4. .5 mm. 

Gall. Small, dark brown, globular galls in clusters of ten to twelve 
on tcnninal twigs of Qucrciis lobala, S mm. in diameter, moi'c or less 
irregular in shape due to compression, the exposed surface rugose. 
Monothalamous, the small, round larval cell, 2 mm. in diameter, 
imbedded centrally in spongy tissue. 

Habitat. Palo Alto, Cal. (Miss Bertha Wiltz). Kern 
County, California. (Beutenmuller.) 

Beutenmiiller describes the gall of this species as "covered 
with a dense, short and compact woolly substance and hairs." 
He states also that it occurs on leaves. His specimens are 
obviously different from mine, although specimens of the fly 
frorn my collection which he has compared with those in his 
possession, he states are similar. 

Cynips maculipennis (Gillette). 

Holcaspis maculipennis, Gillette, Can .Ent., vol. 26 (1894), p. 236. 
Disholcaspis maculipennis, Dalla Torre and Kicffer, Das Tierreich, lief. 24 
Cynipidse (1910), p. 375. 

Female. Rufous, antenna; and abdomen mixed with black, densely 
clothed with sordid white silky pubescence except on abdomen dorsally. 
Head broad, bulging beyond the eyes, faintly punctate, antennte 14- 
segmented, filiform, 1st and 2nd segments stout, 3rd longest, succeed- 
ing segments to 10th progressively shorter, 10th and following seg- 
ments subequal except the last, which is a trifle longer than penulti- 
mate. Pronotum narrow in the middle, mesonotum finely rugose, 
parapsidal grooves deep and reaching to anterior margin, median 
longitudinal lines extending half-way to posterior margin, smooth 
lines over base of wings distinct and rather broad, scutellum finely 
rugose, cushion-shaped, foveje indistinct. Abdomen well developed, 
broader than long, slightly compressed, valves black. Wings large 
and rather broad, hyaline, veins brown with brownish cloud at base of 
cubital cell, another beneath areolet, and numerous small brown spots 
in cubital cell. Length 4.5 mm. 

1911] CynipidcE of California 345 

Gall. Large, globular galls on the leaves of Qucrcus garryana and 
of other species of oak, yellowish brown and covered with rather large, 
dark brown spots, about 3(J mm. in diameter. The outer shell is very 
thin and the single larval chamber is held in place by radiating fibers. 

Habitat. McConaughy (Siskiyou Co.), Cal. (R. W. 

Cynips heldae n. sp. 

Female. Very similar to C. mullipunctala, from which it can 
scarcely be separated, except on the character of the gall. 

Gall (PI. XXIIl, fig. 2). A small gall, with very irregular shape, 
occurring on Quercus lobata; more or less cubical, with many ridged and 
pointed projections, about 8 mm. long and 5 mm. square. ]VI>' spec- 
imens, which are old and dry, are brownish, but in its natural state the 
gall was probably reddish, and is composed of a crystalline substance 
similar to the material fonning the gall of Diplolepis echina. 

Habitat. Ukiah, Cal. (Miss Held). 

Cynips kelloggi n. sp. 

Female. Reddish brown, the eyes, ocelli, a spot on front of head, 
tips of mandibles, oral margin, prosternum, antennse distally from 7th 
segment, metathoracic carinae, abdomen dorsally at ape.K and dorsal 
valve, tibia of hind legs and tarsi black or blackish. Head loigoso- 
punctate, antennaj 15-segmented, 1st segment obconic, 2nd oval, ord 
long, as long as 1 and 2 together, following segments to 10th progres- 
sively shorter, 11th to loth subequal. Thorax rugose, pubescent, 
parapsidal and median grooves complete, median longitudinal lioes 
reaching half-way to posterior margin, smooth lines over base of wings 
distinct, pleura sinooth, shining, punctate in the middle, pubescent 
above and below, seutellum coarsely rugoso-punctate, pubescent, fovece 
rather large, oval, oblique, shallow, with smooth bottom, not approx- 
imate, metanotum rugose. Abdomen smooth and shining, 2nd segment 
occupying about one-half its length, 3rd segment wide but 4th and oth 
narrow, all the segments laterally pubescent, sheaths of ovipositor 
projecting and with dorsal val-ve pubescent. Legs slightly pubescent, 
claws unidentate. Wings hyaline, pubescent, veins black or blackish, 
radial cell open at the margin, its basal vein arcuate, almost angulate, 
areolet distinct, cubital vein reaching basal. Length 2-3 mm. 

Gall. An elongated swelling of the twig of Quercus douglasi, about 
20 mm. long and 10 mm. in diameter, its outer covering the same as the 
bark of the twig. Polythalamous. 

Habitat. Stevens Creek, beyond Cupertino, Cal. (R. W. 
Patterson) . 

346 Annals Entomological Society of America [Vol. IV, 

7 ANDRICUS Hartig. 

Cynips (part.), Linne, Syst. Nat., ed. 10 (1758), p. 553. 
Andricus (part.), T. Hartig, Zeits. f. Ent., vol. 2 (1840), p. 185, 190. 
Andricus, Aphilothrix, A. Forstcr, Verh. Zool. Ges. Wien, vol. 19 Abh. (1869), 
p. 331, 335, 336. 

Andricus, G. Mayr., Gen. d. Cynip. (1881), p. 12. 

Cheeks at most only half the length of the eyes. Antennte of female 
12-16 segmented, of male 14-17 segmented. Pronotimi narrow in the 
middle. Mesonotnm shagreened or nearly smooth, sometimes trans- 
versely folded. Parapsidal grooves usually percurrent. Scutellum 
with two basal foveas, without median line on disc. Metanotal ridges 
parallel or arcuate. Radial cell elongate, open at the margin. Tarsal 
claws bidentate. AlDdomen almost glabrous. Againic and sexual 

Andricus quercus-califomicus (Bass.) 

Cynips quercu,s californica, H. F. Bassc-tt, Can. Ent., vol. 13 (1S81) p. 51. 
Andricus calitornicus, G. Mayr, Gen. d. Cynip. (1881), p. 28. 
Andricus (Callirhytis) califomicus, Ashmead, Tr. Am. Ent. Soc, vol. 12 
(1885), p. 294. 

Female. Reddish brown, eyes, ocelh, tips of mandibles, 3rd to 6th 
and 1st antennal segments proximall}' and ventral valve black or 
blackish. Head, thorax and legs covered with yellowish white pubes- 
cence. Head faintly rugose, broad and bulging laterally beyond the 
eyes, antennae 14-segmented, filiform, 1st and 2nd segments stout, 3rd 
long, longer than 1 and 2 together, succeeding segments to 9th progres- 
sively shorter, 9th and following segments subequal except the last, 
which is twice as long as penultimate. Pronotum narrow in the middle, 
faintly nigose, mesonotum shallowly punctate, parapsidal grooves 
incomplete, reaching slightly beyond the middle, median longitudinal 
lines extending half-way to posterior margin, smooth lines over base of 
wings long and thin, a median bare spot on pleura smooth and shining, 
scutellum cushion-shaped, rugose, with rather indistinct basal foveas. 
Abdomen broad, smooth and shining, dorsally ridged at apex, 2nd seg- 
ment occupying about half its length, ptibescent at sides basally, 
ventral valve and sheath of ovipositor also pubescent. Wings hyaline, 
pubescent, veins brownish, radial cell rather short and open at the 
margin, vein at base angulatc, arcolct large, cubitus almost reaching 
basal vein. Length .5 mm. 

Gall. The familiar "oak-apple;" large, smooth, yellowish white, 
globular galls found on the branches of Quercus lohala, .")0 to 60 mms. or 
more in diamater, sessile, polythalamous, the numerous larval cells 
imbedded internally in a rather dense cellular tissue. 

Habitat. Santa Rosa, Cal. (Miss Josephine Van Wormer). 

Andricus chrysolepidis Ashm. 

Andricus chrysolepidis, Ashmead, Proc. U. S. Nat. Mus., vol. 19 (1896), p. 119. 

"Gall. A very hard, ovate, or globular gall, with a nipple at apex 
and a centrally imbedded larval cell; externally it is covered with a 
dense, fine, short pubescence like the pubescence on a peach, although 
sometimes this is rubbed off. Diameter, 5-8 mm. 

1911] Cynipidce of California 347 

"Gallfly. Female. Length, 3 to 5 mm. Reddish browai, anteniiEe 
and legs bro\vnish yellow, eyes and abdomen dark, reddish brown. 
Head and thorax closely punctate, sparsely pubescent. Antennas 
14-jointed, very slightly thickened at tips. The thorax, besides the 
two parapsidal grooves, which are obsolete anteriorly, has amedian groove 
extending anteriorly for more than half length of the mesonotum, two 
short median grooves anteriorly on each side of this, and the usual 
groove on the shoulders. Scutellum cushion-shaped, rugose, the 
foves- distinct, pleura finch', minutely rugose, slightly striated at base. 
Abdomen polished, the short apical segments under a high power 
show a fine, delicate punctuation,' while the ventral valve projects but 
slightly. Wings glossy, hyaline, veins yellowish, areolet small; neither 
the apex of the submarginal nor the radial vein reach the margin. 

"Types No. 30G6, U. S. N. M. 

" One female, reared from a gall found on Quercus chrysolepis, at 
Colfax, Placer County, California, October 8, 1SS5, by Mr. Albert 
Koebele; and two specimens reared January IS and 29, 1886, from 
same galls. Other of the galls are niunbered 381G U. S. N. M." (W. H. 

I have not seen examples of this species. 

Andricus congregatus Ashm. 

Andricus congregatus, Ashmead, Proc. U. S. Nat. Mus., vol. 19 (1896), p. 120. 

"Gall. An irregular, rugose, yellowish brown woody swelHng, 
containing numerous cells, growing apparently from the extreme tips 
of very slender twigs of Quercus chrysolepis, the gall appearing to have 
a long peduncle, or it may be at the apex of the petal of a leaf, the leaf 
in consequence being aborted. The gall is more or less contracted in 
the middle and varies in length from 2-4 cm. and in diameter from 
1-2 cm. 

" Gallfly. Female. Length, 2 mm. Pale brown or brownish yellow, 
the eight terminal antennal joints, the middle and posterior tibiae, 
metathorax, abdomen dorsally, and wing veins brown. Head and 
thorax closely, unifonnly punctate. Antennas 14 or 1.5-jointed, depend- 
ing upon whether the terminal joint, which presents a rather distinct 
suture, is counted as one or two joints. The terminal joints all appear 
delicately fluted. Mesonotum has three distinct grooves, extending its 
whole length, and the groove on the shoulder is long. Scutellum 
minutely rugose, the foves oblique, distinct, but rather widely sep- 
arated. Wings hyaline, with short pubescence. 

"Type. No. 3068 U. S. N. M. 

"Seven female specimens, received from Prof. E. W. Hilgard, 
Oakland, California, and reared November 10, 1876. The gall also 
occurs on Quercus agrifolia, and Prof. Riley says "a woody deformation 
of the stamniate aments and quite abundant on some trees." (W. H. 
Ashmead) . 

I have not seen examples of this species. 

348 Annals Entomological Society of America [Vol. IV, 

Andricus crystallinus Bass. 

Andricus crystallinus, H. F. Bassett, Tr. Am. Ent. Soc, vol. 26 (1900), p. 319. 

Female. Cherry red to reddish brown, eyes, oral margin, spot on 
front above the insertion of antennas, antennae distally from 7th seg- 
ment and a band on segments 1, 3, 4 and 5, dorsal and subdorsal ^^tta^ 
on mesonotum concurrent with median longitudinal lines and lines 
over base of wings, scutellum at base, metathorax largely, pedicel, 
abdomen dorsally at base and on .3rd, 4th and 5th segments, pecten, 
hind coxas, tips of tarsi and dorsal valve black or blackish. Head 
minutely rugoso-punctate, face pubescent, antennae 14-segmented, 1st 
segment obconic, 2nd oval, 3rd as long as 1 and 2 together, a trifle 
longer than 4th or 5th, succeeding segments to 10th progressively 
shorter, following segments subequal. Pronotum narrow in the middle, 
punctate, mesonotum also punctate, parapsidal grooves complete, 
median longitudinal lines reaching half-way to posterior margin, smooth 
lines over base of wings long and rather broad, pleura smooth and shin- 
ing, pubescent on upper and lower margins, scutellum rugose, fovea? large, 
oval, deep, bottom smooth and shining, approximate, covered throughout 
with dense pubescence. Abdomen smooth and shining, 2nd segment 
occupying about four-fifths of its length, basally pubescent at sides, 3rd 
segment rather wide, 4th, 5th and 6th quite narrow, these segments 
minutely punctate, ovipositor sheaths exserted and pubescent, legs 
light brown, claws bidentate. Wings hyaline, pubescent, subcostal, 
radial, basal and anal veins heavy and brown, others rather faint, radial 
cell open at margin and rather long, its basal vein arcuate, arcolet 
indistinct, cubitus not reaching basal vein. Length about 2.25 mm. 

Gall. Irregularly shaped palisadal galls, in clusters of 5 or 6, on under 
side of leaves of Quercus dumosa, Qucrcns douglasi and Quercus agrifolia? 
and binding leaves together. Outwardly fuzzy. The individual gall is 
about 7 inm. long and 2 to 3 mm. across. 

Habitat. Jasper Ridge, in the vicinity of Stanford Uni- 
versity, Cal. (R. W. Patterson). St. Helena, Cal. (Miss Julia 
Begley). Napa, Cal. (Bassett.) 

Andricus pacificus Ashm. 

Andricus pacificus, Ashmead, Proc. U. S. Nat. Mus., vol. 19 (1896), p. 118. 

Female. Reddish brown, eyes, ocelH, tips of mandibles, and tip of 
dorsal \'alve blackish, 1st abdominal segment more or less fuscous, legs 
and antenna? basally a yellowish brown. Head faintly rugose, 
pubescent, antenna? 14-segmented, 1st and 2nd segments stout, .3rd long, 
longer thna 4th, succeeding segments to 10th progressively shorter, 
10th and following segments subequal except last, which is a trifle 
longer than penultimate. Thorax faintly rugose, parapsidal and median 
grooves on mesonotum reaching half-way to anterior margin, median 
longitudinal lines extending half-way to posterior margin, smooth lines 
over base of wings distinct and long, pleura aciculate, scutelhnn deeply 
rugose, especially at apex, and pubescent, fovea? large, subcircular, 
shallow, with shining, punctate bottom, and contiguous. Abdomen 

1911] Cynipidcc of California 349 

about equal in length to head and thorax together, 2nd segment exten- 
sive and succeeding segments, which are microscopically punctate, 
telescoped, occupying only a fourth its length, valves conspicuous. 
Wings glassy hyaline, pubescent, radial cell open at the margin, areolet 
large, cubitus not reaching basal vein, vein at base of radial cell arcuate. 
Length 2.. 5-3. 5 mm. 

Gall. Dark greenish or ycllowisli brown, drupe-like galls, 2 cm. in 
length and 1 cm. in width, arising from the leaf-buds of Quercus 
chrysolepis, pointed at apex and with more or less obvious nipple, 
surface rugose. Apparently monothalamous. The exit hole of the 
mature insect is at the base. Internally the gall is of a hard, pithy 
structure, and an elongated, cylindrical canal leads to the large cen- 
trally imbedded larval cell. These galls are described by Ashmead as 
smooth and sometimes polished. 

Habitat. Stevens Creek, above Cupertino, Cal. (R. W. 
Patterson.) Placer County, California. (Ashmead.) 

Andricus dasydactyli Ashm. 

Andricus dasydactyli, Ashmead, Proc. U. S. Nat. Mus., vol. 19 (1896), p. 117. 

"Gall. This gall, in structure, is ver\r peculiar, and consists of an 
oblong or elongated, woody tube, in shape not unlike a date seed; it is 
two centimeters long by from one-half to three-fourths of a centimeter 
in diameter, one end being attached sessilly to the branch and covered 
with long, brownish yellow wool. Internally there is a cylindrical 
hollow, which, however, does not extend its entire length, being inter- 
rupted or stopped up by the small larval cell which is situated near 
its center. 

"Sometimes three or more of these galls occur close together on the 
branch, and with their woolly covering present a curious appearance. 
One of the specimens in the collection is almost globular, but all the 
others are as described above. 

"Gall-fi}'. Female. Length 3. S mm. Clear reddish brown ; vertex 
of head and the extreme tip of abdomen dusky; antennas and legs 
brownish yellow. Head and thorax minutely, fineh' punctate, the 
pleura with fine stria?. Antennte 11-jointed, rather long, the 3rd joint 
one-third longer than 4th, the follo\ving to 8th gradually shortening, 
beyond this about equal, the terminal joint being slightly lengthened. 
The parapsidal grooves are only distinct on the posterior half of the 
mesonotum, entirely wanting anteriorly; anteriorly extending to about 
the middle of the mesonotum are two median, glabrous lines ; posteriorly 
there is a long median grooved line, while the line on the shoulders is 
distinct; the scutellum is more coarsely rugose at the apex, the basal 
fovea; large, ovate, oblique, approximate, glabrous at bottom. The 
abdomen is slightly longer than the head and thorax together and of 
the usual shape. Wings glassy hyaline, only shghtly pubescent, the 
veins pale yellowish, except the basal nervure and the vein at base of 
marginal cell, which are brown; this last vein is arcuate but not angulate. 
The areolet is large and the cubital cell is not quite closed. 

"Type. No. 3063, U. S. N. M. 

350 Annals Entomological Society of America [Vol. IV, 

"Described from many female specimens, which issued at various 
dates between January 18, 1SS5 and February 11, 1SS6. The gall 
occurs in California on Quercus chrysolepis, and was sent to the Depart- 
ment of Agriculture by Mr. Albert Koebele." (W. H. Ashmead). 

I have only seen specimens of the gall, which were collected 
from Quercus chrysolepis, in the Stevens Creek Canyon, above 
Cupertino, Cal. It is very characteristic and unmistakable. 

Andricus kingi Bass. 
Andricus kingi, H. F. Bassett, Tr. Am. Ent. Soc, vol. 26 (1900). p. 31G, 317. 

Female. Brown, eyes, ocelli, tips of mandibles, tips of tarsi and 
dorsal valve black. Head finely reticulate, face a ]jale brown, without 
pubescence, antenna; 14-segmented, 1st and 2nd segments stout, 3rd 
segment long, nearly as long as 1 and 2 together, one-third longer than 
4th, 4th and 5th subcqual, succeeding segments to 9th progressively 
shorter, 9th and following segments subcqual, all the segments more or 
less fuscous. Thorax finely reticulate and punctate, covered with a 
sparse pubescence, parapisdal grooves complete and deep, median 
longitudinal lines and smooth lines over base of wings rather indistinct, 
median longitudinal groove from posterior margin very indistinct, 
mesopleura smooth, shining, microscopically reticulate, pubescent 
below, scutellum rugose, basally with large, circular foveae, shallow, 
with smooth bottom, and contiguous, separated only by a carina. 
Abdomen smooth, shining, compressed, 2nd segment occupying about 
one-half its length, following segments wholh^ visible, dorsal valve and 
ovipositor sheaths prominent, last segment reticulate, 3rd, 4th, 5th, 6th 
and margin of 2nd punctate. Wings hyaline, pubescent, with faint 
iridescence, radial cell open and rather long, areolet small, cubitus not 
reaching basal vein. Length 2mm. 

Gall. Small, pink, cone-shaped galls on the under side of leaves of 
Quercus lobata, about 5mm. high and 4 mm. across the base, which is 
broad and rather saucer-shaped. The outer portio^i of the gall is 
conical, the sides curved inwards slightly. The larval chamber is near 
the apex, where the exit hole is found. Some of the galls, from which 
flies were bred in every way identical with those from typical specimens, 
are said to be whitish striped with red and on leaves of Quercus douglasi, 
but specimens are not at hand. 

Andricus parmula Bass. 

Andricus parmula, H. F. Bassett, Tr. Am. Ent. Soc, vol. 26 (1900), p. 312. 

Female. Ferruginotis, the legs and antennce yellowish brown, 
eyes, ocelli, tips of mandibles, tips of tarsi, abdomen dorsally at apex 
and ventral valve black. Head faintly rugose, face pubescent, antennje 
13-segmented, 1st and 2nd segments stout, 3rd segment long, only a 
little longer than 4th, 4th and 5th segments subcqual, succeeding seg- 
ments to 10th progressively shorter, 10th and following segments 
subequal, except last, which is twice as long as pentdtimate. Pronotum 
narrow in the middle, mesonotum coarsely reticulate, parapsidal grooves 

1911] Cynipidce of California 351 

incomplete, reaching but half-way to anterior margin, median longi- 
tudinal lines extending half-way to posterior margin, smooth lines over 
base of wings distinct, pleura finely striated, with a triangular smooth 
area posteriorly, scutellum rugose, sparsely pubescent, basal foveas 
large, elliptic, oblique, with smooth bottom, not approximate, metano- 
tal ridges only slightly curved, faced with black. Abdomen about as 
long as head and thorax together, smooth and shining, more or less 
lenticular in shape, second segment occupying half its length, 3rd 
segment rather broad, following segments narrow, ovipositor sheath 
exserted, concolorous. Wings rather opaque whitish, faintly iridescent, 
veins faint, yellowish, radial cell narrow, open at the margin, vein at 
base arcuate, almost angulate, areolet indistinct, cubitus not reaching 
basal vein. Length 2 mm. 

Gall. Very small, flat, reddish, disc-shaped galls on the under side 
of leaves of Quercus lobata, about 3 mm. in diameter, slightly elevated in 
the center. Much smaller than gall of Andricus pailersonm and quite 
distinct from it. 

Andricus wisliceni Ashm. 
Andricus wisliceni, Ashmead, Proc. U. S. Nat. Mus., vol. 19 (lS9o), p. 119. 

"Gall. A small, globular gall, with a slight projection at base 
where it is attached to the twig ; it varies in color from a 3'ellow brown to 
dark brown, and some are mottled with purple and brown. It is hard, 
and contains in the center a small larval cell; diameter, 3-4 mm. 

"Gall-fly. Female. Length, 3 to 3.4 mm. Pale brownish j-ellow, 
almost devoid of pubescence, the abdomen polished and di.scolored witn 
brown, eyes dark brown, the mandibles black. Head and thorax finely 
punctate, shining; in front of the anterior ocellus is a deep transverse 
fovete, and there are some coarse scattered punctures on the mesonotum. 
Antenna: 14-jointed, the 3rd joint about one-eighth longer than the 4th, 
the joints from 7th to apex short, about twice as long as wide, dusky, 
and delicately fluted. Parapsidal grooves distinct, the groove on the 
shoulders very long, distinct, and a little bent anteriorly. Scutellum 
cushion-shaped, rugose, the foveas at base large and distinct, separated 
only by a slight carina ; pleura smooth, but under a high power showing 
faint delicate striae. Wings hyaline, the pubescence short, veins, 
except the sub-marginal vein from the portion extending from the 
basal vein to apex, and the angulated cross vein at base of marginal cell, 
which are brownish or piceous, yellowish. 

"Type. No. 3065, U. S. N. M. 

"Nine female specimens, reared October 14, 1886, from the galls 
sent to the National Museum by Mr. Albert Koebele, collected in 
Sacramento County, California, on Quercus wisliceni." (W.H.A.shmead). 

I have not seen examples of this species. 

352 An7ials Entomological Society of America [Vol. IV, 

Andricus quercus-flocci (Walsh). 

? Cynips quercus lana, A. Fitch, 5th Rep. In.s. N. York (1859), p. 34. 

? Cvnips quercus lanae, Osten Sackcn, Proc. Ent. Soc. Philad., vol. 1 (1861), 
p. 62. 

Cvnips quercus llocci, B. D. Walsh, Proc. Ent. Soc. Philad., vol. 2 (1864), 
p. 482: 

Cvnips (Andricus) flocci Ostcn Sacken, Proc. Ent. Soc. Philad., vol. 4 (1865), 
p. Zai. 

Andricus flocci, G. Mavr, Gen. d, Cvnip. (1881), p. 28. 

? Andricus lana, Ashmead, Tr. Am. Ent. Soc, vol. 12 (1885), p. 295. 

Female. Black, ocelli, antenna;, legs distally from the coxas, 
teguke and ovipositor .sheath brownish. Head reticulately nigose, 
shining, face wth whitish pulx'scenee, antennas lo-segmented, 3rd 
segment longest, not as long as 1 and 2 together and only a little longer 
than 4th, following segments progressi-\-ely shorter except last, which is 
one and one-half times the length of penultimate, distal segments 
fuscous. Pronotum narrow in the middle, rugose and pubescent, 
mesonotum smooth and shining, or more or less shagreened, ]jarapsidal 
grooves deeply impressed, complete, the median longitudinal lines 
reaching more than half-way to posterior margin, smooth lines over 
base of wings distinct and reaching anterior margin, pleura aciculate, 
seutellum rugose, slightly jjubesccnt, basal fovea; large, shallow, with 
smooth bottom, approximate. Abdomen smooth, shining, greatly 
compressed apieally, 2nd segment oecujn'ing more than half its length, 
pubescent at base, 3rd and following segments naiTow, ventral valve 
perpendicular, reaching tergal line, ovipositor exserted and curving 
upward, ovipositor sheath pale, pubescent. Wings hyaline, pubescent, 
veins brown, radial cell long, open at the margin, vein at base arcuate, 
areolet small, cubitus not reaching basal vein. Length 2.2,i mm. 

Gall. A rnass of twenty or more smooth, brown, elliptical or loaf- 
shaped galls, 2mm. high and 1 mm. in diameter, on under side of leaves 
of Quercus lobata, covered with reddish or yellowish wool. Alonothal- 

Habitat. Palo Alto, Cal. (Miss Bertha Wiltz.) 

Andricus patter sonae n. sj:). 

Female. Very similar to A. kirigi. from which it can scarcely be 
separated, except on the character of the gall. Some of the specimens 
have the abdomen dorsally and the antenna: distally from 9th segment 
more or less blackish, and dorsal and subdorsal vittjc on mesonotum 
concurrent with the median longitttdinal lines and smooth lines over 
base of wings. 

Gall (PL XXIII, fig. 3). Thin, flat, disc-shaped galls from leaves 
(presumably) of Quercus douglasi, about (i min. in diameter, greenish 
gray with lilac center, the margin irregular; upper surface a trifle 
wrinkled but otherwise smooth. 

Habitat. Stanford Univer.sity, Cal. (R. W. Patterson.) 

1911] Cynipidce of California 353 

Andricus wiltzffi n. sp. 

Female. Head and thorax black, abdomen walnut brown, darker 
(almost black) on the dorsum, antennae bro^vn, 1st, 3rd and 4th segments 
spotted with black, distally from 6th segnemt fuscous to black, tegulag, 
sheath of ovipositor, anterior legs and joints of middle and hind legs 
brown. Head broad and thin, with close, reticular sculpturing, eyes 
narrow, antennas i;]-segmcnted, 1st and 2nd segments stout, 1st obconic, 
2nd subglobular, 3rd longest, as long as 1 and 2 together, following 
segments progressi\'cly sliorter to last, which is twice penultimate. 
Prothorax narrow in the middle and transversely folded, sparsely 
punctate, each puncture bearing a hair, mesonotum opcnh^ reticulate, 
transversely folded, parapsidal grooves complete and distinct, median 
longitudinal lines reaching almost half-way to posterior margin, smooth 
lines over base of wings distinct, pleura smooth and polished, aciculate 
in the middle, pubescent below, scutellum deeply rugose, basal iovei^ 
large, oval, with smooth, shining bottom, approximate, separated only 
by a carina, metanotum smooth, polished, pubescent. Abdomen stout, 
subglobose, smooth and shining, 2nd segment occupj'ing less than 
one-half its length, 3rd segment rather broad, .slightly punctate, 4th, 
5th and (ith segments narrow, all with oblique margins, ventral valve 
concealed, black, ovipositor sheath slightly projecting, pubescent. 
Wings hyaline, pubescent, subcostal, basal and radial veins heavy, 
brown, others faint, radial cell long, narrow, open at the margin, vein 
at base arcuate, areolet distinct but veins on two sides faint, cubitus 
faint, not reaching basal vein. Length 2. .5 mm. 

Gall. Polythalamous galls formed in the buds of Qucrciis lobata, 
distinguished by the thicklj- compacted cluster of aborted leaves. 

Habitat. Stanford University, Cal. (Miss Bertha Wiltz.) 

Andricus brunneus n. sp. 
Female. Brown to reddish brown, the eyes, ocelli, tips of mandibles, 
antennte distally, metanotum in the middle, tips of tarsi, abdomen 
dorsally at apex and dorsal valve black or blackish. Head faintly 
reticulate, face pubescent antennae 14-segmented, 1st segment obconic, 
2nd oval, ord longest, as long as 1 and 2 together and a little longer than 
4th, succeeding segments to 9th progressively shorter, 9th and following 
segments subeciual, except the last, which is a little longer than jDenulti- 
mate. Pronotum and mesonotum finely reticulate, sparsely punctate, 
each puncture bearing a hair, parapsidal grooves indistinct, reaching 
but half-way to anterior margin and rather widely sejaarated, median 
longitudinal lines extending half-way to posterior margin, smooth lines 
over base of wings distinct and long, pleura smooth, microscopically 
reticulate, pubescent abo\-e and below, sctitellum rugose, basally with 
large, transverse fovese, smooth at bottom, contiguous, separated only 
by a carina, metanotum almost smooth. Abdomen smooth, shining, 
compressed and dorsallj' ridged, 2nd segment occupying not more than 
one-third its length, pubescent at the base, 3rd, 4th, 5th and Gth segments 
punctate, 7th segment reticulate, dorsal valve and sheaths of ovipositor 

354 Annals Entomological Society of America [Vol. IV, 

exserted. Wings hyaline, pubescent, subcostal, basal and vein at base 
of radial cell heavy, brown, others faint, radial cell long, open at margin, 
vein at base arcuate, with fuscous suffusion, areolet small but distinct, 
cubitus not reaching basal vein. Length 3 mm. 

Gall. Thin-shelled, subglobular galls on leaves of Quercus douglasi, 
about the size of a pea, pointed at opposite poles. 

Habitat. Stanford University, Cal. (R. W. Patterson.) 


Callirhytis, A. Forster, Verb. Ziol. Ges. Wien, vol. 19 Abh. (1869), p. 331, 

Andricus (Callirhytis), G. Mayr, Gen. d. Cynip. (1861), p. 27. 

Differs from Andricus only in the following particulars: 
parapsidal grooves not always complete, tarsal claws simple. 

Callirhytis chrysolepidicola (Ashm.) 
Cynips chrysolepidicola, Ashmead, Proc. U. S. Nat. Mus., vol. 19 (1896), 
p. 124. 

Female. Browai, eyes, tips of mandibles, face above base of antennas 
and on anterior margin, and metathoracic carinje black, antennae, 
abdomen dorsally, and tibiae and tarsi of middle and hind legs fuscous. 
Head rugose, slightly pubescent, antennas 15-segmented, 1st and 2nd 
segments stout, 1st obconic, 2nd smaller, 3rd segment longest, longer 
than 1 and 2 together, 4th-6th segments progressively shorter, following 
segments subequal except the last, which is smaller than penultimate. 
Mesothorax punctate and pubescent, parapsidal grooves indistinct, 
scutcllum small, cushion-shaped, rugose and pubescent, fovea? distinct, 
oval, shallow and approximate. Abdomen darker than head or thorax, 
smooth and shining, 2nd segment pubescent at base, ventral and dorsal 
valves of medium length and pubescent, ovipositor long, extending 
much beyond the abdomen. Wings hyaline, pubescent, subcostal, radial 
basal and 2nd transverse veins heav\-, black, other veins rather feeble, 
radial cell open at margin, areolet distinct. Length 2 mm. 

Gall. Stem or twig galls; brown, with a bluish tint, and rugose, 
over 2.5 mm. long and about 15-20 inms. in diameter, the long axis lying 
in the direction of the twig, hard and woody, polythalamous. 

Habitat. Pacific Grove, Cal. (C. P. Smith.) Pine Canyon, 
Cal. (Ashmead.) 

Callirhytis apicalis (Ashm.) 

Andricus apicalis, Ashmead. Proc. U. S. Xat. Mus., vol. 19 (1896), p. 120. 
Callirhytis apicalis, G. Mayr, Verh. Zool. Ges. Wien, vol. 52 (1902), p. 289. 

"Galls. Irregtdar, brownish black, globular galls of a dense pithy 
substance, growing on the roots of Quercus wisliceni, sometimes three 
or four together, pressing each other into irregular shapes. Diameter 
usually about half an inch. 

"(jall-fly. Female. Length 5.8 to 7 mm. Bright brick red, the 
mandibles black at tips. Head and thorax finely punctate with some 
larger, coarser ptmctttres scattered over the surface, and almost free 

1911] CynipidcE of California 355 

from pubescence. Checks full, bulging. Antennae 14-jointed, filiform, 
the 3rd joint slightly larger than 4th and narrowed toward base, the 
apical joint twice as long as the preceding, fusiform. Parapsidal 
grooves distinct, a more or less distinct medial groove and distinct 
grooved lines on the shoulders. Scutellum rugose, with two large 
foveffi at base, separated by a carina; pleura anteriorly slightly rugose, 
posteriorly nearly smooth, with some very delicate striae. Abdomen 
smooth, with a few hairs on the side of second segment; the tenninal 
segments show a fine, delicate punctuation; the spine of the ventral 
valve is long. Wings hyaline, except the entire apical third, which is 
smok}^ or dark brown, the veins stout, black, the angular projection in 
marginal cell at base being vfirv distinct. 

"Type. No. -.3067, U. S. N. M. 

"Three specimens, reared by Mr. Albert Koebele, fomi galls collected 
in Sacramento County, California, but the year of collecting and the 
date of rearing are not given. A single specimen (No. 3714) was reared 
February 17, 1SS6. The bright red color and smoky bases of wings will 
readily distinguish the species." (W. H. Ashmead). 

I have not seen examples of this species. 

Callirhytis quercus-pomiformis (Bass.) 

Cynips quercus pomiformis, H. F. Bas.sett, Can. Ent., vol. 13 (1881) p. 74. 

Andricus pomiformis, Ashmead, Tr. Am. Ent. Soc, vol. 12 (188.5), p. 29.5. 

Callirhytis pomiformis, G. Mayr, V'erh. Zool. Ges. Wien, vol. 52 (1902), 
p. 289. 

Callirhytis quercus-pomiformis, Dalla Torre and Kieffer, Das Tierreich, lief. 
24 Cynipidce (1910), p. 568. 

Female. Black, antennas, legs, mandibles and abdomen ventrally 
dark brown. Head coarsely rugose, antennte 14-1.5 seginented, 1st and 
2nd segments stout, 3rd segment longest, a little longer than 4th, suc- 
ceeding segments to 9th progressively shorter, 9th and following seg- 
ments subequal, 15th a mere tip on preceding segment. Thora.K 
coarsely rugose, parapsidal grooves distinct, complete, median longi- 
tudinal lines reaching half-way to posterior margin, sctitellum rugose, 
with subcircular basal fovea" shallow, smooth and shining at bottom, 
not exactly approximate. Abdomen more or less smooth and shining, 
2nd segment occupying about two-thirds its length, ventral vah'e and 
sheath of ovipositor pubescent. Legs clothed with pubescence. Wings 
hyaline, radial cell open at the margin, a black cloud at base, subcostal 
vein scarcely reaching margin, vein at base of radial cell angtdate, areolet 
distinct, cubitus not always reaching basal vein. Length 3 min. 

Gall. Yellowish, subsphcrical galls, about 3.5 mm. in diameter, 
the surface more or less roughened by slight elevations in the fonn of 
longitudinal ridges, containing within numerous oval larval cells im- 
bedded in the pithy substance of the gall. 

Habitat. Stanford University, Cal. (R. W. Patterson.) 

356 Annals Entomological Society of America [Vol. IV, 

Callirhytis quercus-agrifoliae (Bass.) 
Cvnips quercus agrifoliip, H. F. BassLtt, Can. Ent. vol. 13 (1881), p. 53. 
Andricus (Callirhytis) agrifoliae, G. Mayr, Gen. d. Cynip. (1881), p. 28. 
Callirhytis quercus-agritolia;, Dalla Torre and Kieffer, Das Tierreich, lief. 
24 Cynipida; (1910), p. 567. 

Female. Pale yellowish brown, the antennsc distally from lOth 
segment and abdomen dorsally fuscous to black, ej^es, ocelli, tips of man- 
dibles, prosterntim, metathorax, tips of tarsi and ventral valve black. 
Head faintly rugose, almost granulose, face pubescent, antenna? 14-seg- 
mented, filiform, 1st and 2nd segments stout, 1st obconic, 2nd o^'al, .3rd 
segment longest, as long as 1 and 2 together and a little longer than 4th, 
succeeding segments to 9th progressively .shorter, 9th-13th subequal, last 
segment longer than penultimate by a half. Thorax elevated, subspher- 
ical, pronotum narrow in the middle, granulose, slightly pubescent, 
mesonotum reticulate, almost shagreened, parapsidal grooves deeply 
impressed, not quite complete but approaching anterior margin, median 
longitudinal lines reaching half-way to posterior margin, smooth lines 
over base of -^vings long and very distinct, pleura largely granulose, 
slighth^ pubescent above and beneath, aciculate in the middle, scutellum 
granulose, pubescent, basal foveae large, oval, transverse, and shallow, 
with smooth, shining bottom, approximate, metathorax shallowly 
punctate, pubescent. Abdomen long oval, smooth and shining, 2nd 
segment occupying about half its length, 3rd and 4th segments rather 
wide, succeeding segments narrow, posterior margins in all oblique, 
oviposito'r sheath exserted, pubescent. Wings hyaline, pubescent, 
veins pale brown, radial cell long, narrow, open at the margin, vein at 
base angulate, areolet distinct, cubitus not reaching basal vein. 
Length 3. .5 mm. 

Gall. Moderately large, brownish, hard, globular galls, about 
10 mms. in diameter, on twigs of Quercus agrifolia, surface microscopic- 
all}^ pubescent. Internally composed of a dense, cork-like substance, 
in which the single larval cell is imbedded. 

Habitat. Palo Alto, Cal. (R. W. Patterson.) 

Callirhytis lasia Ashm. 

Callirhytis lasius, Ashmead, Proc. U. S. Nat. Mus., vol. 19 (1896), p. 132. 

Female. Pale brownish yellow, legs Hghter than body, abdomen 
dorsally darker, head and thorax pubescent. Head faintly rugose, 
antennae 14-segmented, Lst segments obconic, 2nd long oval, .3rd seginent 
longest, as long as 1 and 2 together and incised proximally for two-thirds 
its length, succeeding segments to 9th progressively shorter, 9th and 
following segments, which are fuscous, subequal, except last, which is 
longer than penultimate. Thorax closely punctate, punctation rather 
coarse, parapsidal and median grooves from posterior margin distinct, 
the former almost complete, median longitudinal lines reaching nearly 
half-way to posterior margin, smooth lines over base of wings rather 
long, scutellum flat, closely punctate, basal foveas transverse and not 
separated by a carina. Abdomen smooth and shining, 2nd segment 
oceup\-ing less than half its length, pubescent at base, 3rd, 4th and .5th 

1911] Cynipidce of California 357 

tergitcs visible, dorsal and ventral valves with a brush of hairs. Wings 
hyaline, pubscent, veins farily distinct, radial cell open at margin, short, 
vein at base arcuate, radius only slightly bent, almost a straight line, 
areolet present and distinct, cubitus not reaching basal vein. Length 
2 mm. 

Gall. Brownish, subglobular, depressed galls on under side of 
leaves of Qucrcus chrysolepis, (1-7 mm. in diameter, punctate, and covered 
with pubescence, which is pale brownish tinged with red. Polythal- 

Habitat. Stevens Creek Canyon, above Cupertino, Cal. 
(R. W. Patterson.) 

Callirhytis quercus-suttoni (Bass.) 

Cvnips quercus suttoni, H. F. Bassett, Can. Ent., vol. 13 (1881), p. 54. 
Aridricus (Callirhytis) suttoni, G. Mayr, Gen. d. Cynip. (1881), p. 28. 
Callirhytis quercus-suttoni, Dalla Torre and Kieffer, Das Ticrreich, lief. 
24 CynipidEe (1910), p. 564. 

Callirhytis polythyra, Beutenniuller in litt. 

Female. Reddish brown, tips of mandibles, oral margin, antenna; 
distally from 4th segment at joints, prostemtmi, base of scutellum, 
metathorax, abdomen dorsally at base, ventral valve and hind tibite 
and tarsi black or blackish. Head faintly rugose, face pubescent, eyes 
narrow, antennae 15-segmented, filifonn, 1st and 2nd segments stout, 
.3rd segment longest, longer than 1 and 2 together and one-half longer 
than 4th, sticeeeding segments to 9th progressively shorter, 9th-l,5th 
subequal. Pronotum narrow in the middle, mesonotum coriaceous 
and sparsely punctate, pubescent, parapsidal and median grooves 
reaching less than half-way to anterior margin, median longitudinal 
lines and smooth lines over base of wings distinct, pleura pubescent, 
medially smooth and bare, scutellum rugose and pubescent, a rather 
broad, arcuate groove at base in place of the usual fovea;. Abdomen 
smooth and shining, second segment occupjang less thaii half its length, 
pubescent at base, 3rd and 4th segments rather broad, succeeding 
segments narrow or concealed, ventral valve prominent, ovipositor 
sheath exserted, pubescent. Legs thickly covered with short hairs. 
Wings hyaline, pubescent, veins brownish, rather faint, radial cell long, 
narrow, open at the margin, vein at base angulate, areolet distinct, 
cubitus not reaching basal vein. Length 3.5 mm. 

Gall. Spherical or elongate twig swelling on Quercus agrifolia, and 
Qucrcus chrysolepis, about 20-30 mms. in length and 1.5-25 mms. in 
diameter, covered with normal bark and ligneous within. Polythal- 
amous, the numerous oval cells lying near the surface. 

Habitat. Claremont, Cal. (Baker.) 

Callirhytis vacciniifoliae Ashm. 

Callirhytis vacciniifolia;, Ashmead, Proc. U. S. Nat. Mus., vol. 19 (1896), 
p. 130. 

"Gall. A thin shelled, globtilar gall, with a central kernel held in 
place by radiating filaments and closely resembling the gall produced 

358 Annals Entomological Society of America [Vol, IV, 

by A. inanis Osten-Sacken, but the average size is smaller and the color 
of the gall darker. It measures from half an inch to a little over an 
inch in diameter and is found on Quercus vacciniifolia in California. 

"Gall-fly. Female. Length, 2 to 3.2 mm. Red or brownish 
red (one specimen has the thorax almost black), antenna; and legs 
brownish yellow, sometimes obfuscated; several terminal joints of the 
antennae are dark brown. Head closely punctate ; thorax almost smooth, 
the parapsidal grooves sharply defined, complete, a short inedian groove 
posteriorly and the groove on the shoulder indistinct ; scutellum rugose, 
the fovea: large, broad, distinct, and separated only by a carina; meso- 
pleura smooth, polished, the triangular piece beneath tegulas alone being 
punctate. The abdomen is longer than the head and thorax together, 
the segments oblique, the second segment occupies about two-thirds 
of the whole surface and is a little pubescent at sides near the base, 
impunctate, some of the short terminal segments a little dusky; spine 
of ventral valve rather long, hairy. Wings hyaline, pubescent, the 
veins pale brown; the vein at base of marginal cell is slightly bent, but 
not angulate, the marginal cell being very long and narrow, areolet 
distinct, but the surrounding veins delicate, cubital cell almost closed. 

"One of the specimens is but 2 mm. long and of a uniform brownish 
yellow color, but structurally it does not seem to differ from the others. 

"Type No. 3087, U. S. N. M. 

"Four specimens, reared October 16, and December 4, 1884, from 
galls collected by Prof. J. H. Comstock, at Truckee, California, on 
Quercus vacciniifolia." (W. H. Ashmead). 

I have not seen examples of this species. 

Callirhytis maculipennis Kieff. 

Callirhytis maculipennis, Kicffer, Bull. Soc. Metz, st-r. 2, vol. H (1904), 
p. 131. 

Callirhytis maculipennis, Kieffer, Invert. Pacif., vol. 1 (1904), p. 42. 

"Black, opaque and pubescent. Head rugose, enlarged behind the 
eyes; front coriaceous, temples longitudinally striated and finely punc- 
tured, nearly smooth and shining close behind the eyes. Antennse 
brownish-red, the last joints more or less fuscous, 14-jointed, nearly 
glabrous; second joint hardly longer than thick; the third at least 
thrice as long as thick, the following joints gradually decreasing in 
size; joints 9-13 nearly equal, one third longer than thick; the last a 
little longer than the penultimate. Thorax coarsely rugose; mesonotum 
irregularly rugose posteriorly between the parapsidal furrows, anteriorly 
and laterally more or less transversely rugtilose; parapsidal furrows 
complete; the four glabrous lines of the mesonotum are rigid; meso- 
pleura shagreened and smooth; scutellum hardly longer than wide, 
rugose, with two triangular fovea? at the base and a longitudinal middle 
furrow in the anterior two-thirds. Metanotum glabrous, shagreened, 
with three longitudinal carinae, the lateral ones curved outwards. 
Wings hyaline, fringed, with a fuscous spot at the base of the radial 
cell; first abscissa of the radius angulate. Legs brownish red, hind 
femora and middle part of the hind tibiae fuscous; claws simple. Abdo- 

1911] Cynipidce of California 359 

men smooth, shining, laterally pubescent only at the base of the second 
segment, longer than the head and the thorax united; second segment 
one-fourth shorter than the whole abdomen, finely punctulate apically, 
laterally very sloping; the two following segments also punctulate. 
Length of female 3.5-4 mm. 

"Gall. A bud gall on Querciis agrifolia, globular, the diameter 
22 mm. or more, whitish, smooth and glabrous, apically wath a very 
small wart; the inner substance is also whitish, somewhat spongiose, 
and contains below the middle several bro^vnish cells which are 4 mm. 
high and 2 mm. wide. 

"This species is allied to the genus Amphibolips, from which it is 
excluded by the tarsal claws being simple. " (J. J. Kieffer). 

I have examined an example of both gall and gallfly of C. 
maculipennis. kindly furnished by Mr. Baker. 

Callirhytis eriophora Kieff. 

Callirhytis eriophora, Kieffer, Bull. See. Metz, ser. 2, vol. 11 (1904), p. 132. 
Callirhytis eriophora, Kieffer, Invert. Pacif., vol. 1 (1904), p. 43. 

"Brownish red, opaque, scarcely pubescent, and finely shagreened. 
Head enlarged behind the eyes. Antennte 14-jointed, the second joint 
longer than thick ; the third fully thrice as long as thick, scarcely longer 
than the fourth ; the following joints gradually shorter and thicker ; the 
penidtimate, one-third longer than thick, shorter than last. Mesono- 
tum nearly glabrous, with four dusky longitudinal stripes, the lateral 
of which are abbreviated anteriorly, the two others shortened posteriorly ; 
parapsidal furrows not complete, anteriorly obliterated. Scutellum 
rugose, nearly glabrous, basally with two divergent fovea. Metanotum 
black-brown, with two parallel carina. Wings hyaline, fringed, the 
veins brown, the first abscissa of radius angulatcd. Legs faint yellow; 
claws simple. Abdomen dark red-brown, above nearly black; the 
second segment occupying nearly the whole abdomen, very shining, 
glabrous, posteriorly microscopically punctulate; ventral-spine eight 
times as long as wide. Length of the female 3-3.2 mm. 

" Gall. A bud gall on Quercus ii'isUzeuii; the gall is roimded, 10 mm. 
in diameter, the outer portion a yellowish wool-like substance, the inner 
a globular, ligneous, thin-shelled and monothalamous cell, 4 mm. or 
more in diameter." (J. J. Kieffer). 

I have seen examples of this species, kindly furnished by 
Mr. Baker. 

Callirhytis clarimontis Kieff. 

Callirhytis clarimontis, Kieffer, Bull. See. Metz, ser. 2, vol. 11 (1904), p. 132. 
Callirhytis clarimontis, Kieffer, Invert. Pacif., vol. 1 (1904), p. 43. 

"Brownish yellow, finely shagreened and scarcely shining. Head 
enlarged behind the eyes. Antennae black-brown apically, 14-jointed; 
the third joint nearly five times as long as thick, one half longer than 
the fourth; the following ' joints gradually decreasing in length, the 
penultimate one-third longer than thick, the last more than twice as 

360 Annals Entomological Society of America [Vol. IV, 

long as thick. Mesonotum nearly glabrous, with four dusky longi- 
tudinal stripes, the two lateral ones abbreviated anteriorly; the parap- 
sidal furrows complete. Scutellum rugose, nearly glabrous, with two 
divergent fovea? at the base. Metanotum black-brown, with two 
parallel carina?. Wings hyaline, fringed, the veins dusky, the first 
abscissa of the radius angulated. Legs and coxte faint yellow, claws 
simple. Abdomen glabrous, highly polished, not punctulate, dorsally 
more or less black-brown, the second segment surpassing somewhat 
the middle, laterally very sloping; ventral-spine four to five times as 
long as wide. Length of the female S.o-t mm. 

"Gall. The gall of this species was found on Quercus agrifolia; it is 
a bud gall S mm. in diameter and more, globular, opaque, yellowish, 
with some sparse brown spots, glabrous, apically with short and sparse 
hairs and a small wart; the inner substance is brown, spongiosc, and 
encloses a globular, faint yellow, thick-shelled, monothalamous shell 
5 mm. in diameter and more; wall of the cell 1.6 mm. thick.' ' (J.J.Kieffer) 

I have seen examples of this species, kindly furnished by 
Mr. Baker. 

Callirhytis bakeri Kieff. 

Callirhytis bakeri, Kieffc-r, Bull. Soc. Metz, ser. 2, vol. 11 (1904), p. 132. 
Callirhytis bakeri, Kieffer, Invert. Pacif., vol. 1 (1904), p. 44. 

"Brown-red, shagreened and pubescent. Head enlarged behind 
the eyes. Antennae sometimes black-brown apically, 14-jointed; the 
second joint longer than thick; the third joint four times as long as thick, 
one-third longer than the fourth; joints 4-G subequal, the fourth scarcely 
longer than the fifth, the eight apical joints gradually decreasing in 
length, the penultimate one-half longer than thick, a little shorter than 
the last, or sometimes only half the length of the last. Thorax scarcely 
shining ; parapsidal furrows obliterated anteriorly ; scutellum rugose, the 
two basal fovea? black, deep, separated by a small carina and oblique 
and divergent; postscutellum and metanotum rugose, the two carinas 
parallel. Wings hyaline, fringed, veins brown, first abscissa of the 
radius angulated. Legs and coxa? faint yellow, claws simple. Abdomen 
black-brown, highly polished and shining, glabrous, longer than the 
head and the thorax united, the second segment with a pubescent spot 
on either side at base; these spots occupy the basal three-fourths, the 
segments are microscopically punctured apically; ventral-spine four 
times as long as wide. Length of the female .'J.S-4 mm. 

"Gall. A bud gall on Quercus crassipociila [chrysolepis]; gall oval, 
about 13 mm. high and 11 mm. thick, smooth, glabrous, ligneous, 
apically with a thin, longitudinally striated, thick and more or less bent 
point which is about S mm. high and 1..5 mm. thick; proximally with 
some bud-scales lying close to the base of the gall; the inner substance 
subspongiose. " (J. J. Eeffer). 

I have seen examples of this species, kindly furnished by 
Mr. Baker. 

1911] Cynipidce of California 361 

Callirhytis rossi Kieff. 

Callirhytis rossi, Kieffer, Marcellia, vol. 2 (1903), p. 84. 

"Forme Agame. Noir; mandibules, les sept premiers articles 
antennaires, pattes anterieures, tibias et tarses des pattes inter- 
mediaires, et ecaillettes ferrugineux; scape plus ou moins noiratre; 
toutes les hanches noires; femurs intermediaires et posterieurs, tibias et 
tarses des pattes posterieures d'un brun noir. Abdomen d'un brun 
marron, dessus noir en partie. Axrtcx avec des rides irregulieres et 
grossieres; face avec des stries rayonnant de la bouche jusqu' au bord 
des yeux; joues egalant la moitie de la longeur de ces derniers. 
Mandibules bidentees, aussi larges que longues. Palpes maxillaires 
composes de quatre articles, dent le premier depasse de moitie la 
longeur du second; cclui-ci a peine plus long que le troisieme, trois fois 
aussi long que gros; le dernier un peu plus long que le premier et dis- 
tinctcment plus gros. Palpes labiaux comjjoses de trois articles dont 
le second n' est pas plus long que gros; le premier cylindrique, presque 
trois fois aussi long que gros; le troisieme jslus gros que les autres, un 
peu aminci aux deux bouts, avec une petite nodosity a I'extremite, 
deux foix aussi long que gros, a peu pres aussi long que le premier. 
Antennes a, peine plus epaisses vers I'extremite, composees de 14 articles, 
dont le second est deux fois aussi long que gros; Ic troisieme quatre fois 
aussi long que gros, un peu plus long que le quatriemc ; les suivants 
diminuant graduellement ; le douzieme et le treizieme a j^eine plus 
longs que gros, le quatorzieme double du treizieme. Thorax mat, a 
peine pubescent; mesopleures avec une tache brillants et lisse; partie 
infericure des propleures ridec regulierement ct longitudinalement; 
pronotum, mesonotum, scutellum et segment mediaire rides grossiere- 
ment et irregulierement ; sillons parapsidaux percurrents, entre eux sc 
voient deux aretes paralleles situees dans le tiers anterieur du mesono- 
tum; c6tes du mesonotum avec un sillon longitudinal bien apparent. 
Fossettes du scutellum profondes, tr^s rapprochees k leur base. Seg- 
ment mediaire avec deux aretes faiblement eonvergentes et circon- 
scrivant une aire traversec par une arete en forme de T; cotes du 
segment mediaire avec une dent distinete. Crochets des tarses simples, 
a piene aussi longs que la palette. Ailes anterieures faiblement 
enfum(Ses dans leur tiers apical, avec une tache d'un brun marron a 
Tangle basal interne de la memc cellule; ncrvnire sous-scotalc marquee 
de brun noir a I'endroit de I'insertion de la basale; bord cilie; cellule 
radiale peu longue, ouverte a la merge, largement aussi a la base et 
faiblement au sommet; premiere partie du radius arquee, au moins 
aussi longue que la partie apicale de la sous-costale ; areole distinete; 
nervures d'un brun elair, base du cubitus peu marquee. Ailes infer- 
ieures hyalines, avec une nervure sous-costale, une basale s'etendant 
des crochets frenaux au bord posterieur et une mediane s'arretant a la 
basale. Abdomen avec une large tache de pubescence de chaque cote 
de sa base, avec sept segments distincts, aussi long que la tete et le 
thorax r^unis; second segment occupant la moitie de la longeur de 
I'abdomen; bord posterieur des six premiers segments finement et 

362 Annals Entomological Society of America [Vol. IV, 

•densement pointille; apres la mort, a moins que les insectes n'aient ete 
tues dans I'alcool, le second segment atteint presque I'extreniite abdom- 
inal, et ne laisse a decouvert qu'un mince bord des deux segments 
suivants. Spinule ventrale trois fois aussi longue que large. Taille 
9 4^5 mm. 

"Oeuf presque globuleux, un peu plus long que gros, atteignant la 
cinquieme de la longueur de son pedicclle. 

"Galle. On trouve la galle de cet insecte sur un Chene a feuilles 
caduques et legerement pubescent sur le dessous, a jeunes rameaux 
tomenteux de blanc. [Quercus chrysolepis?] Elle est formee aux 
depens d'un bourgeon axillaire; comme celle de Cynips tozce Bosc, elle 
chevauche sur le rameau qui, pour I'unique excmplaire que j'ai recu, 
etait gros de 3 mm. Sa forme est spherique, sa couleur d'un jaune 
brunatre, son diametre de 40 mm., sa surfcae couverte d'asperites 
surtout dans la moitie superieure; ces asperites atteignent au maximum 
1 mm. en hauteur et sont souvent confluentes de facon a former des 
•carenes ou des rides irregulieres, longitudinales ou transversales. Au 
pole superieur, la galle se prolonge en une pointe coniquc, haute de 5 
mm., large d'autant a sa base, ligneuse, et striee longitudinalement. 
L'interieur est spongieux comme chez celle de Cynips tozce, avec le 
centre lignieux; cctte partie ligneuse et tres dure, dont le diametre 
mesure IS mm., renferme de nombreuses cellules larv^aires separees 
I'une de I'autre par une parol epaisse de 0.60 a 1 mm., longues de 3.5 a 
4 mm. et larges de 2 mm. Chacune de ces cellules renferme une galle 
interne de meme forme, etroitement appliquee au tissu environment, 
mais d'un gris jaunatre un peu plus clair, se detachant assez facilement 
et a parol extrement mince et fragile. Eclosion en mai de la seconde 
annee. J'ai obtenu 19 Cynipides d'une seule galle. 

"Patrie. Cahfomie. L'unique exemplaire m'a ete cnvoye par 
M. le docteur Ross, a qui I'insecte est dedie.'' (J. J. Kieffer.) 

I have not seen examples of this species. 

Callirhytis nigra n. sp. 
Female. Black, ocelli, antenna', tcgulae, legs and ovipositor sheath 
brown. Head broad and thin, microscopically reticulate, face and 
cheeks pubescent, antennae 13-segmented, incrassate at tip, 1st and 
2nd segments stout, 2nd oval, about one-half of 1st, 3rd and 1st seg- 
ments subequal, 4th-6th a little shorter, 7th and following segments 
shorter and subequal, except the last, which is twice as long as penulti- 
mate and a little longer than 3rd; all the segments more or less pubes- 
cent. Pronotum narrow in the middle, sides deeply punctate, each 
puncture bearing a hair, mesonotum coriaceo-reticulate, almost smooth 
and shining, parapsidal grooves distinct and complete, median longi- 
tudinal lines not reaching half-way to posterior margin, smooth lines 
over base of wings distinct, mesopleura smooth and shining, aciculate 
above, pubescent beneath, scutellum deeply rugose, basal foveas oval, 
deep, ^\'ith smooth, shining bottom, approximate. Abdomen smooth 
and shining, much compressed apically, 2nd segment occupj'ing about 
half its length, its posterior margin oblique, 3rd and 4th segments nar- 

1911] Cynipidce of California 363 

row, ventral valve oblique and extending upwards to dorsal line, ovi- 
positor sheath protruding slightly and pubescent. Legs covered with 
pubescence. Wings hyaline, pubescent, veins faint, brownish, radial 
cell long, open at the margin, vein at base arcuate, areolet and cubitus 
indistinct. Length 1.75 mm. 

Gall. An irregular swelling or enlargement of the terminal twigs 
of Qucrciis lobata, subglobular, 1.5 mm. x 12 mm., apparently 
mono thalamous . 

Habitat. . San Jose, Cal. (R. W. Patterson.) 

Callirhytis guadaloupensis n. sp. 

Female. Yellowish brown to reddish brown, one specimen with the 
head, pronotum, median stripe on mesonotum, scutellum, metanotum 
and abdomen darker; tips of mandibles, ocelli, basal region of anten- 
nae, pccten, tibiae, tarsi and caudal asjiect of femora black or blackish. 
Head closelj' punctate, pubescent, antennas 14-segmented, 1st and 2nd 
segments stout, 3rd segment long, longer than 1 and 2 together, 4th to 
8th segments progressively shorter, following segments subequal, 
except last, which is twice penultimate. Thorax rugose and partially 
pubescent, ]}arapsidal grooves on mesonotum reaching half-way to 
anterior margin, median longitudinal lines and smooth lines over base 
of wings distinct, the former extending less than half-way to posterior 
margin, scutelhmi pointed, deeply rugose and pubescent, basal foveas 
inconspicuous, pleura rugose and pubescent. Abdomen smooth, six 
tergites visible, the second occupying less than one-third the entire 
length, pubescent at base, dorsal and ventral valves pubescent, tips 
with tufts of hair. Wings small and narrow, veins heavy, outer third 
of cubitus not reaching margin, the basal portion of which is hairy, 
areolet distinct, basal third of cubitus not reaching basal vein. 
Length 2.5 mm. 

Gall (PI. XXIII, fig. 4). Hard, smooth, flat, disc-shaped galls on 
under side of leaves of Querciis chrysolepis, 5-6 mm. in diameter 
apparently monothalamous. 

Habitat. Guadaloupe, Cal. (R. W. Patterson.) 

Callirhytis sanctae-clarae n. sp. 

Female. Reddish brown, eyes, tips of mandibles and ocelli black, 
legs yellowish brown. Head faintly rugoso-punetate, pubescent, 
antennse 14-segmented, 1st and 2nd segments stout, 3rd segment long, 
a little longer than 1 and 2 together, succeeding segments to 9th pro- 
gressively shorter, 9th and following segments subequal, except the 
last, which is a little longer than penultimate. Thorax rugoso- 
punetate, parapsidal grooves on mesonotum distinct, extending half- 
way to anterior margin, median longitudinal lines reaching half-way to 
posterior margin, smooth lines over base of wings distinct, mesopleura 
almost bare, upper portion closely and finely punctate, lower portion 
acieulate, scutellum rugoso-punetate, basal foveas large, shallow, with 
smooth, shining bottom, approximate, separated only by a carina, 

364 Annals Entomological Society of America [Vol. IV, 

metanotum pubescent. Abdomen smooth, shining, very much com- 
pressed, 2nd segment occupying more than two-thirds its length, 
posterior margin finely punctate, ventral valve conspicuous, paler than 
the rest of the abdomen, dorsal valve darker and pubescent, ovipositor 
sheath conspicuously exserted. Wings hyaline, pubescent, veins dis- 
tinct, yellowish, radial cell, open at the margin, vein at base arcuate, 
almost angulate, areolet distinct, cubitus not reaching basal vein. 
Length 3-3.5 mm. 

Gall (PI. XXIII, fig. 5). Acorn galls at the tip of leafy twigs of 
Qiiercus cbrysolepis. 15 mms. long, and double, the basal portion 
smaller, and telescoped in the upper portion; rugose and covered with a 
whitish bloom. Monothalamous. 

Habitat. Stevens Creek Canyon, above Cupertino, Cal. 
(R. W. Patterson.) 

9 SYNERGUS Hartig. 

Synergus (part.), T. Hartig, Zeits. f. Ent., vol. 2 (1840), p. 186, 197. 

Face radiately striated, clypeus imperceptible or nearly so, subdorsal 
frontal grooves always present, usually long. Antenna? 13-15 seg- 
mented in female, 14-15 segmented in male, distally from 3rd segment in 
female filiform, in male often thickened, 3rd segment often strongly emar- 
ginate basally and distally more or less thick. Mesonotum shagreened, 
very often transversely wrinkled, parapsidal grooves complete, scutellum 
with basal fove^. Abdomen compressed, 1st segment or petiole longi- 
tudinally striate, tergite of the 2nd segment greatly enlarged and 
covering the greater part of the abdomen, or in the male obscuring it, 
2nd and 3rd tergites joined without suture and enclosing following 
segments. Radial cell closed, short. Claws bidentate. Ventral valve 
short. Inquilines in galls on Qiiercus. 

Synergus agrifolise Ashm. 

Synergus agrifolise, Ashniead, Tr. Am. Ent. See, vol. 2.3 (1896), p. 189. 

"Female. Length 2-2.4 mm. yellow, the legs and coxae 
paler, the metathorax dusky; ocelli, eyes and mandibles black, or brown- 
black. The face and pleura are finely striated, head and thorax finely 
. minutely rugose, pubescent, the parapsidal furrows wanting, the foveas 
of scutellum small, shallow, hardly apparent; antennas 13-jointed, 
flagellar joints 2 and 3 equal, little shorter than the first. The abdomen 
is not quite as long as the head and thorax combined, slightly conpressed, 
and vertically it is almost as wide as long with the tips sometimes 
dusky. Wings hyaline, pubescent, the veins pale or hyaline, the areolet 
distinct, but the closing vein very delicate; in two specimens it is absent. 

"The male is but 1.5 mm. long, and, excepting the black eyes and 
ocelli, entirely brownish yellow. 

"Habitat. Los Angeles, California. 

"Described from thirteen specimens, bred by Mr. Albert Koebele, 
from a gall not unlike Neiiroterus saltatoriiis Edw., occurring on Qiiercus 
agrifolia." (W. H. Ashmead). 

I have not seen examples of this species. 

1911] CynipidcE of Califoniia 365 

Synergus brevicomis Ashm. 

Sj-nergus brevicomis, Ashmcad, Tr. Am. Ent. Soc, vol. 23 (1S96), p. ISO. 

"Female. Length 2.<S-3 mm. Stature similar to S. laviventris, 
but the sculpture of the thorax is more rugose, the abdomen proportion- 
ately shorter, the ventral valve projects slightly, and the antennje reach 
back only two-thirds the length of thorax. Vertex of head, thorax and 
abdomen black; orbits, cheeks, face, antenna- and legs, brownish yellow,, 
the depth of color often varying; middle and posterior coxae black, the 
front jjair often dusky basally, posterior femora often more or less 
clouded; antennas 18-jointed, filifonn, the 3rd joint one-third longer than 
4th; face and mesopleura rather coarselj^ striated; scutellum rounded, 
depressed at base with two distinct foveae ; the grooves of the mesonotum 
rounded, almost obsolete by the coarse sculpture. Abdomen highly 
polished, the second segment entirely hiding all the others and gaping 
open at apex, in consequence the ventral ^^alvc projects, is brownish 
and armed with a slight spine. Wings hyaline, pubescent, the veins 
pale, the submarginal and marginal veins brownish apically; areolet 
large, closed, the cul^ital cell only partially closed. 

"The male is but l.S mm. long, with the head wholly, except stem- 
maticum, and the legs, including coxae, brownish yellow. The antennae 
are short, 15-jointed, the 3rd joint long, strongly excised, the apical 
joint brown. 

"Habitat. Eldorado County, California." (W. H. Ashinead). 

I have not seen examples of this species. 

Synergus flavus Kieff. 
Synergus flavus, Kieffer, Bull. Soc. Metz, ser. 2, vol. U (1904), p. 1.33. 
Synergus flavus, Kieffer, Invert. Pacif., vol. 1 (,1'J04), p. 45. 

"Pale yellow or reddish yellow; head distinctly broader than the 
thorax, without carina; front and vertex smooth, face and cheeks paler 
yellow. Antennas of the male 15-jointed, the two to three last joints 
more dusky; second joint longer than thick; the third scarcely longer 
than the fourth, faintly emargined outwardly; the fourth twice as long 
as thick ; the following ones gradually becoming shorter ; the penultimate 
joint is a little longer than thick, shorter than the last and all are very 
slender; antennas of the feinale 14-jointed, filiform and slender; the third 
joint fully thrice as long as thick, the following gradually decreasing 
in size, the penultimate only one-third longer than thick, shorter than 
the last. Mesonotum finely coriaceous; mesopleura very finely longi- 
tudinally striated; scutellum brownish red, rugose, the usual foveas small, 
not very distinct, separated by a small carina. Metanotum brown or 
black-brown, the carinee parallel. Wings hyaline, veins brownish. 
Tarsal claws simple. Petiole of abdomen black, coarsely longitudinally 
striated; the second segment reaching to the apex of the abdomen, 
smooth and not punctured. ■• 

"Bred from galls of CalUrhytis maculipeunls; the guests live in the 
spongiose substance of the gall, outside of the cells, and do not hinder 
the development of the gall makers. " (J.J. Kieff erj. 

I have seen examples of this species, kindly furnished by 
Mr. Baker. 

366 Annals Entomological Society of America [Vol. lY, 

Synergus dimorphus O.-S. 

Svnergus dimorphus, Osten-Sacken, Proc. Ent. Soc. Philad., vol. 4 (1865), 
p. 376. 

Synergus dimorphus, Gillette, Tr. Am. Ent. Soc, vol. 23 (1896), p. 86. 87. 

Female. Black, antennas, ocelli, face on anterior margin, mandibles 
basally, gense and orbital margins, tegulas, legs (except coxas) and 
ovipositor sheath yellowish brown, tips of antennae and tarsi fuscous. 
Head faintly reticulate, face radiately striate, antennas 14-segmented, 
1st segment stout, 2nd slender and oval, 3rd longest, slightlji- longer 
than either 4 and 5, following segments progressively shorter, except 
the last, which is somewhat longer than penultimate. Pronotum wide 
in the middle, rugose, mesonotum transversely wrinkled (some speci- 
mens without transverse folds, rugose), parapsidal grooves incomplete, 
extending less than half-way to anterior margin, median longitudinal 
lines not reaching the middle, smooth lines over base of wings distinct, 
mesopleura transversely aciculate, with a triangular smooth patch 
beneath wdng, scutellum deeply rugose, fovese indistinct, small, oval and 
approximate, separated by a median carina; pronotum, mesonotum and 
scutellum sparsely covered with silvery pubescence. Abdomen smooth 
and shining, vertical and horizontal dimensions almost equal, tergal line 
from the side almost straight, sternal line semicircular, 2nd tcrgite 
wholly obscuring the following segments, posterior margin faintly 
punctate. Wings hyaline, pubescent, radial, subcostal, basal and 2nd 
transverse veins black, heavy, others faint, radial cell short, closed, vein 
at base slightly arcuate, areolet indistinct, cubitus not reaching basal 
vein. Length 2.25-2.5 mms. 

Male. Black, tips of mandibles, palpi, antenna (except distally 
infuscated basal segments), legs (except trochanters and coxas), ventral 
margin of abdomen and a transverse basal band brown. Head rugose 
and finely reticulate, antennte 15-segmented, reaching middle of abdo- 
men, 3rd and 4th segments subequal, 3rd longer than 1 and 2 together, 
incised at base, following segments progressively shorter to last. Thorax 
coarsely rugose, mesonotum transversely wrinkled, parapsidal grooves 
complete and deeply impressed, median longitudinal lines distinct, 
posterior margin of abdomen punctate. Wings hyaline, pubescent, 
areolet distinct, cubitus nearly reaching basal vein. Length 3.5 mm. 

Bred from galls of Cynips multipunclata on Quercus lobata. 

Habitat. Stevens Creek Canyon, above Cupertino, Cal. 
(R. W. Patterson.) 

Synergus oneratus oneratus (W. Harr.) 

Cynips oneratus, T. W. Harris, Treat. Ins. N. Eng. (1842), p. 398. 

Synergus oneratus, B. D. Walsh, Proc. Ent. Soc. Philad., vol. 2 (1864), p. 
488, 498. 

Synergus oneratus oneratus, Dalla Torre and Kieflfer, Das Tierreich, lief. 
24 Cynipidaj (1910), p. 632. 

Female. Yellowish brown, legs and face light yellowish brown, 
eyes, ocelli, tips of mandibles, prosternum, pecten, mesopleura beneath, 
metathorax apically and hind tarsi black; jjubescent. Head faintly 

1911] Cyuipida of California 367 

rugose and sparsely punctate, punctation in the form of irregularly 
scattered shallow pits, face pubescent, radiately striate, antennae 
14-segmented, 1st segment large, 2nd subglobular, 3rd and 4th segments 
subequal, as long as 1 and 2 together, 5th and succeeding segments to 
11th progressively shorter, 11th to 14th subequal. Pronotum wide in 
the middle, rugose, mesonotum trans\'crsely wrinkled, parapsidal 
grooves distinct, extending half-way to anterior margin, median longi- 
tudinal lines not reaching the middle, mesopleura coarsely aciculate, 
with a triangular smooth patch beneath wings, scutellum coarsely 
rugose, almost coarsely reticulate, basal fovers transversely elongate, 
shallow, smooth and shining at bottom, approximate, metanotum 
rugose. Abdomen smooth and shining, 2nd tergite completely obscur- 
ing the following segments, posterior margin microscopically punctate, 
apex dorsally black, dorsal valve prominent, pubescent, ovipositor 
protruding. Wings hyaline, pubescent, a few of the veins heavy and 
black, the others faint, radial cell closed, very short, vein at base not 
much curved, areolct small and indistinct, veins enclosing it alirtost 
obliterated, cubitus not reaching basal vein and very faint. Length 
2-3 mm. 

Male. Black, antennae, face beneath antenna: and around eyes, 
ocelli, tegulas, coxee, trochanters, femora, tibia; largely, and abdomen on 
posterior margin brownish or pale brown; sparsely pubescent. Head 
reticulate, faintly ptmctate, face radiately striate, antennae 15-seg- 
mented, all the segments conspicuously grooved or channeled, 3rd and 
4th segments subequal, longer than 1 and 2 together, succeeding seg- 
ments to 11th progressively shorter, 11th and following segments 
subequal. Thorax coarsely rugose, mesonotuna tranversely ridged, 
parapsidal grooves indistinct, almost wholly obliterated by the coarse- 
ness of sculpture, pleura coarsely aciculate, small median patch smooth, 
scutellum coarsely rugose, almost coarsely reticulate, smooth and shin- 
ing beneath ruga;, fovete large, oval and shallow, with smooth bottoms, 
separated only by a carina, metanotum rugose. Abdomen smooth and 
shining, 2nd tergite completely obscuring foUomng segments, posterior 
margin microscopically punctate. Wings hyaline, pubescent, costal, 
subcostal, radial and basal veins heavy, black, others faint, radial cell 
closed, areolet small, indistinct, cubital vein faint and not reaching 
basal, vein at base of radial cell almost straight. Length 1.5-2 mm. 

Bred from galls of Ilolocaspis eldoradensis on Quercus lohata. 
Habitat. Jasper Ridge, near Stanford University, Cal. 
(R. W. Patterson.) 

Synergus punctatus Gillette. 
Synergus punctatus, Gillette, Tr. Am. Ent. Soc, vol. 23 (1896), p. 90, 94. 

Female. Black, the ocelli, face anteriorly, antennae, tegulae, pedicel, 
abdomen anteriorly and posteriorly and legs brown ; sparsely pubescent. 
Head faintly reticulate, shining, face radiately striate, antenna 14-seg- 
mented, 1st and 2nd segments stout, 3rd long, about as long as 1 and 2 
together and longer than 4th, succeeding segments to 10th progressively 

368 Annals Entotnological Society of America [Vol. IV, 

shorter, 10th and follo-n-ing segments snbequal, except last, which is 
longer than the penultimate, all longitudinally grooved or channeled 
and pubescent. Pronotum Isroad in the middle, rugose, mesonotum 
transversely wrinkled, folds rather shallow, parapsidal grooves reeaching 
less than half-way to anterior margin, median longitudinal lines and 
smooth lines over base of \'^-ings distinct, mesopleura aciculate, triangu- 
lar patch beneath wings smooth and shining, scutellum deeply rugose, 
fovccB large, circular shallow, metathorax almost smooth. Abdomen 
smooth and shining, 2nd tergite completely obscuring following seg- 
ments, pubescent at base, posterior margin punctate, sheath of ovipos- 
itor conspicuous, pubescent. Wings hyaline, ])ubescent, veins basally 
heavy, black, radial cell closed, short, vein at base only slightly bent, 
areolet subobsolete, onl}'' one of its enclosing veins distinct, cubitus not 
reaching basal vein. Length 1.25 mm. 

Bred from galls of Ilolcaspis eldoradensis on Quercus lobata. 
Habitat. Jasper Ridge, near Stanford University, Cal. 
(R. W. Patterson.) 

Synergus ochreus n. sp. 

Female. Brown, the legs and face light brown, eyes, ocelli, tips of 
mandibles, pecten, dorsal vitta on abdomen, ventral valve and tips of 
tarsi black. Head rugose, face radiately striate, subdorsal strife from 
base of antennae to ocelli and continued around eyes, antennas 1.5-seg- 
mented, 1st segment stout, obconic, 2nd narrowly ovate, half the length 
of the 1st, 3rd-6th subequal, one-third longer than 1st, 7th-9th progres- 
sively shorter, 10th and following segments subequal. Pronotum wide 
in the middle, trans\'erscl3' rugose, pubescent, mesonotum transversel}' 
wrinkled or folded, parapsidal grooves complete, widely separated, 
obscured by transverse folds, a median longitudinal groove posteriorly 
reaches half-way to anterior margin, median longitudinal lines short, 
smooth lines over base of wings distinct, mesopleura transversely 
aciculate and shining, pubescent above and below, scutellum cushion- 
shaped, rounded posteriorl}^ faintly rugose and punctate, each puncture 
bearing a hair, basal foveas large, circular or transversely oval and 
shallow, bottom punctate, shining, pubescent. Abdomen compressed, 
smooth and shining, 2nd tergite reaching apex and almost wholly 
obscuring following segments, faintly but broadly punctate on posterior 
margin, ventral \'ah-e reaching line of the tergum, black. Wings 
hyaline, pubescent, veins, excepting anal and cubital, heavy, brown, 
radial cell short, closed, ^■ein at base slightly arcuate, areolet ab.sent, 
veins on two sides and basal abscissa of cubitus scaixely visible. 
Length 3.5 mm. 

Bred from galls of Cynips miiJti punctata on Quercus lobata. 
Habitat. Stevens- Creek Canyon, above Cupertino, Cal. 
(R. W. Patterson.) 

1911] Cy^iipidce of California 369 

Synergus niger n. sp. 

Female. Black, antenna;, ocelli, tips of mandibles, tegulfe, legs 
(except hind coxs basally), abdomen on ventral and posterior margin, 
and ovipositor brown, last segment of antennse fuscous. Head broad, 
smooth and shining, microscopically reticulate, face radiately striate 
except in a circular smooth patch above mouth, antenna 14-segmented, 
rather stout, ,1st and 2nd segments stouter than those following, 3rd 
segment longest, longer than 4th but not as long as 1 and 2 together, 
following segments progressively shorter, except last, which is much 
longer than ]jenultimate. Thorax smooth and shining, • pronotum 
broad in the middle, microscopically coriaceo-reticulate, pubescent, 
mesonotum finely sculptured and punctate, each puncture bearing a 
hair, parapsidal grooves incomplete, reaching about the middle, median 
longitudinal groove coextensive, median longitudinal lines extending 
less than half-way to posterior margin, smooth lines over base of wings 
distinct, mesopleura smooth and shining, transversely aciculate in middle, 
pubescent beneath, scutellum deeply rugose, fove£e large, subcircular 
and shallow, with smooth, shining bottom, contiguous, separated by a 
carina, metanotum smooth, punctate and pubescent. Abdomen 
smooth and shining, 2nd tergite extending to apical margin and almost 
wholly obscuring the following segments, nearly as long as head and 
thorax together, but not wide, line of the tergum only slightly arcuate, 
ovipositor sheath exserted. Wings hyaline, pubescent, veins brown, 
only subcostal, radial and basal nervures distinct, radial cell closed, 
vein at base slightly arcuate, areolet indistinct or absent, cubitus 
indistinct. Length 1.75-2 mms. 

Male. Differs in having 15-segmented antennee and the head 
brown below base of antennae, on cheeks and on orbital margin. 

Bred from a btid gall on Quercus lobata. 

Habitat. Stanford University, Cal. (R. W. Patterson.) 

Synergus splendidus n. sp. 
Female. Reddish brown, the eyes, tips of mandibles, occiput 
dorsally and front almost to base of antennae except a narrow orbital 
margin, antennte basally, pronotum in the middle, pecten, mesopleura 
basally, mesonotum, scutellum, metathorax, 1st abdominal segment 
and dorsal ridge of 2nd tergite basally black. Head broad and thick, 
eyes bulging, microscopically reticulate, face radiately striate, pubescent, 
antenna 15-segmented, 1st and 2nd segments stout, obconic, touched 
with black, 3rd. segment longest, a little longer than 4th but not as long - 
as 1 and 2 together, following segments progressively shorter, the last, 
which is fuscous, not much longer than penultimate. Pronotum wide 
in the middle, rugose and microscopically coriaceo-reticulate, covered 
with short appressed hairs, mesonotum rugose and distinctly trans- 
versely wrinkled, parapsidal grooves complete, median longitudinal 
lines widely separated and not reaching the middle, smooth lines over 
base of wings distinct, mesopleura transversely aciculate, pubescent 
beneath, scutellum rugose, basal foveas small, oval, smooth at bottom, 

370 Annals Entomological Society of America [Vol. IV, 

approximate, separated only by a carina, metathorax smooth and 
covered with short appressed hairs. Abdomen longer than head and 
thorax together, greatly compressed, smooth and shining, 2nd and 3rd 
tergites connate, reaching apical margin, incised dorsally at apex for 
one-third the length, exposing the following segments, posterior margin 
of all the segments minutely punctate, ovipositor sheath projecting 
obliquely from venter and reaching line of the tergum, ovipositor 
exserted. Wings hyaline and clothed with rather long, erect hairs, 
veins distinct and fuscous, radial cell short, closed, vein at base .slightly 
arcuate, areolet small, cubital vein reaching basal. Length 5 mm. 

Found in jar with galls from Quercus lobata. 
Habitat. California. 

Synergus multiplicatus n. sp. 

Female. Black, the ocelli, face anteriorly, genae, orbital margin 
and margin of occiput, antennae, pronotum (except medially), mesopleura 
and legs (except hind tibi;e and tarsi) yellowish to reddish brown. Head 
broad, width twice length, coarsely rugoso-punctate, transversely 
rugose on vertex, antennae 14-segmented, 1st and 2nd segments obconic, 
3rd segment longest, as long as 1 and 2 together and longer than 4th by 
a half, 5th and followang segments progressively shorter, except the last, 
which is nearly twice as long as penultimate, with fuscous tip. Thorax 
rugose, mesonotum transversely wrinkled, parapsidal grooves complete, 
median longitudinal lines and smooth lines over base of wings distinct, 
mesopleura coarsely aciculate, with small triangular patch beneath 
wing smooth, fuscous beneath and jDubesccnt. scutellum coarsely rugose, 
almost coarsely reticulate, fovcEe longitudinal, oblique and shallow, 
bottom rough, not exactly approximate. Abdomen smooth, 2nd and 
3rd tergites connate, covering the greater part of the abdomen, 4th 
tergite narrow posteriorly, all traced with microscopic reticulation, 
dorsal valve and sheath of ovipositor projecting slightly and pubescent. 
Wings hyaline, pubescent, veins brownish to black, radial cell closed, 
vein at base arcuate, areolet small, cubitus not reaching basal vein. 
Length L75-3 mm. 

Male. Similar to female exce]3t legs infuscate, metanotum Ijlack, 
connate 2nd and 3rd tergites obscuring followdng segments, brownish 
on posterior margin and punctate, face yellowish brown instead of 
reddish brown, length about 2 mm. 

Bred from galls of Cynips kelloggi on Quercus douglasi. 

Habitat. Jasper Ridge, near Stanford University, Cal. 
(R. W. Patterson.) Frohm, Cal. (R. W. Patterson.) Paso 
Robles, Cal. (John Morehouse.) San Jose, Cal. (R. W. 

inil] Cyuipida of California 371 

Synergus varicolor n. sp. 

Female. Black, thorax mixed with brown, antennae (except tip),. 
face below antennae, genae, orbital margin broacUy, tegulae, abdomen on 
ventral and posterior margins and legs wholly browai. Head reticulate, 
face radiately striate, slightly pubescent, antennas 14-segiTiented, 3rd 
segment longest, longer than 4th, 4th-(.)th subequal, following seginents 
progressively shorter, except the last, which is a little longer than penul- 
timate. Pronotum wide in the middle, mesonotum rugoso-punctate, 
pubescent, parapsidal grooves incomplete, reaching half-way to anterior 
margin, median longitudinal lines and smooth lines over base of wings 
distinct, mcsopleura aciculate anteriorly, posteriorly smooth and shining, 
scutellum deeply rugose, basal foveas large, oval, and shallow, bottom 
rough, contiguous, separated by a carina. Abdomen highly polished, 
2nd and .'jrd tcrgites connate, wiioUy obscuring following segments, 
line of the tergum only slightly curved, sternal line semicircular, only 
tip of ovipositor, which is brown, protruding. Wings hyaline, pubescent, 
radial cell closed, arcolct and basal part of cubitus rather indistinct. 
Length ^..j mm. 

Male. Differs in having 1 o-segmented antennae and the head 
almost entirely, pro thorax, mesonotum, mesopleura and ventral and 
posterior margin of abdomen brown. 

Bred from galls of Callirhylis pomiformis on Qucrcus agrifolia. 
Habitat. Stanford University, Cal. (R. W. Patterson.) 

Synergus maculatus n. sp. 

Female. Yellowish brown, the eyes, ocelli, tips of mandibles, 
prostenium, metathorax and abdomen dorsally at apex black. Head 
finely punctate, face radiately striate, antennas 14-segmented, 1st and 
2nd segments stout, 3rd segment longest, slightly longer than 4th, 
succeeding segments to 10th progressively shorter, llth-13th subequal, 
last somewhat longer than penultimate. Thorax faintly punctate and 
pubescent, parapsidal grooves incomplete, reaching half-w^ay to anterior 
margin, scutellum rugose, foveas indistinct. Abdomen as long as head 
and thorax together, ellijitical when viewed from above, oval as seen 
from the side, smooth and shining, 2nd and 3rd tergites connate, reach- 
ing apex, and almost completely obscuring following segments, dorsal 
valve and ovipositor sheath conspicuous, slightly protruding. Wings 
hyaline, veins faint, areolet distinct, faint on two sides, radial cell closed. 
Length 1.75 mm. 

The male differs in having stout 15-segmented antennae and more 
black on abdomen dorsally. 

Bred from a small, yellowish-brown, depressed globular 
gall, about 2 mms. in diameter, on upper surface of leaves of 
Qnerciis agrifolia. 

Habitat. Stanford University, Cal. (R. W. Patterson.) 

372' Annals Entomological Society of America [Vol. IV, 

Synergus dubiosus n. sp. 

Female. Yellowish brown, eyes, tips of mandibles, occiput dorsally 
continued on to vertex and front almost to the base of the antennas, 
with the exception of a narrow strip on eye, pronotum in the middle, 
pecten, pleura beneath, mesonotum, scutellum and metathorax entirely, 
pedicel and abdomen dorsally almost to apex black. Head faintly 
rugose, sparsely and shallowly punctate on vertex, face radiately striate, 
pubescent, antenna 14-segmented, thick but filiform, 1st and 2nd seg- 
ments stout, 3rd segment long, as long as 1 and 2 together, 3rd-5th 
subequal, 6th and following segments progressively shorter to last. 
Pronotum faintly rugose, closely punctate, clothed with short, appressed, 
whitish pubescence, parapsidal grooves incomplete, reaching more than 
half-way to anterior margin, median longitudinal lines and smooth 
lines over base of wings distinct, mesopleura transversely aciculate, 
scutellum deeply rugose, basal fove£e large, oval and shallow, bottom 
rough, contiguous, separated by a median carina. Abdomen short, 
not quite as long as the thorax, smooth and shining, 2nd and 3rd tergites 
connate, completely concealing following segments, elliptical from above, 
triangular from side, posterior margin truncate and narrowly punctate. 
Wings hyaline, pubescent, veins black, radial cell closed, areolet distinct, 
cubitus not reaching basal vein. Length 4 mm. 

The male has 15-segmented antennae. 

Bred from galls of CalUrhytis pomiformis on Quercus agrifolia. 
Habitat. Stanford University, Cal. (R. W. Patterson.) 

10 CEROPTRES Hartig. 

Ceroptres, T. Hartig, Zeits. f. Ent., vol. 2 (1840), p. 186, 197. 

Face vnth two parallel ridges from insertion of antennte to clypeus, 
beyond these striate, antenna in female 12-14 segmented, distally 
thickened, in male 15-segmented, sometimes 14-segmented. Parapsidal 
grooves usually not reaching anterior margin of mesonotum, scutellum 
with two basal foveas. 2nd and 3rd tergites more or less connate, 
covering nearly the entire abdomen. Radial cell closed, ventral valve 
short, claws bidentate. Inquilines in galls on Quercus. 

Ceroptres pomiformis Ashm. 
Ceroptres pomiformis, Ashmead, Tr. Am. Ent. Soc, vol. 12 (1885), p. 300. 

"Male. Length .05 inch. Slender, head yellowish brown, with a 
dark brown blotch enclosing ocelli; eyes brown; antennas 15- jointed, 
yellowish, with the apical third brown; thorax black, finely pubescent; 
abdomen bright yellowish brown, infuscated towards tip; wings hyaline, 
veins yellow, radial area closed, narrow. 

"This pretty little species is described from two specimens bred 
from gall of Andricus pomiformis Bass., sent to me from California." 
(W. H. Ashmead). 

I have not seen examples of this species. 

1911] Cynipidce of California 373 

Ceroptres dorsalis Prov. 

Ceroptres dorsalis, Provancher, Addit. Hym. Quebec (1888), p. 398. 

" 9 Long. .18 pee. D'un beau jaune-miel, avec les yeux, une 
tache sur le vertex, le lobe median du mesonotum, Ic metanotum et une 
ligne sur le dos des premiers segments de 1' abdomen, noir. Les mandi- 
bulcs noires a 1' extremite. Les antenncs avec les pattes, jaune sans 
aucune tache. La face fortcment striee et d'un jaune plus pale. Ailes 
hyalines, a nervures brunes, legerement velues, la radiale fcrmee en 
avant. Abdomen comprime, a tariere redress^e, depassant la ligne du 
dos. Los Angeles (Coquillett). 

"Superbe esp^ce, bien remarquable par sa taille. 

" cf Meme coloration que dans la 9 , mais tres remarquable par 
son duexieme segment abdominal qui se developpe en une double 
ecaille depassant I'extrcmite de 1' abdomen et se prolongeant en dessous 
d' aumoinsle double del'epaisseur des autres segments." (L. Provancher.) 

I have not seen examples of this species. 

Ceroptres niger n. sp. 

Female. Black, femora distally, tarsi, oral margin and sheath of 
ovipositor brown; somewhat pubescent. Head faintly reticulate and 
shining, face striate, pubescent, antennas 13-segmented, 3rd segment as 
long as 1 and 2 together, longer than 4th, succeeding segments to 10th 
progressively shorter, 10th and following segments subequal, except the 
last, which is twice as long as penultimate. Pronotum wide in the 
middle, rugose, mesonotum faintly reticulate and shining, parapsidal 
grooves not reaching anterior margin, mesopleura aciculate, a median 
patch smooth and shining, scutellum coarsely rugose, almost coarsely 
reticulate, foveas transverse, large, oval and shallow, with smooth 
bottom, widely separated. Abdomen smooth and shining, 2nd tergite 
reaching more than half-way to apex and pubescent at base, 3rd tergite 
rather wide, 4th and 5th narrow, faintly punctate on posterior margin, 
ovipositor sheath exserted, dorsal valve conspicuous and pubescent. 
Legs clothed with pubescence. Wings hyaline, pubescent, radial cell 
closed, short, basal vein not much curved, areolet present but enclosing 
veins indistinct, cubitus not reaching basal vein. Length 1.5 mm. 

Bred from galls of Holcaspis eldoradensis on Quercus lobata. 
Habitat. Jasper Ridge, near Stanford University, CaL 
(R. W. Patterson.) 


Aylax (part.), T. Hartig, Zeits. f. Ent., vol. 2 (1840), p. 186, 195. 
Periclistus, A. Forster, Verh. Zool. Ges. Wien, vol. 19 Abh. (1869), p. 332, 337. 

Face radiately striate, antennas filiform, in female usually 12-seg- 
mented, in male 14-segmented, mesonotum faintly rugoso-punctate and 
pubescent, parapsidal grooves complete or sometimes not reaching 
anterior margin, scutellum with two transverse basal fovea, 1st abdom- 
inal segment or petiole deeply striate, 2nd and 3rd tergites connate, 
covering a large part of the abdomen. Radial cell short, closed. In- 
quilines in different galls. 

374 Annals Entomological Society of America [Vol. IV, 

Pcriclistus calif omicus Ashm. 

Periclistus califomicus, Ashmead, Tr. Am. Ent. Soc, vol. 23 (1896). p. 188. 

"Male and female. Length 2-2.4 mm. Similar to P. sniihicis. the 
punctuation finer, the pubescence denser, antenna; dark brown. Legs 
reddish yellow, sometimes obfuscated, the middle and posterior coxas 
black, shining. The parapsidal grooves are only distinct on the pos- 
terior half of the mesonotum, entirely wanting anteriorly, and there is 
no distinct grooved line on the shoulders, while the short anterior median 
grooves are wanting; scutellum rugose, foveas large, distinct, oblique; 
mesopleura smooth, highly polished. Wings hyaline, pubescent, 
iridescent, veins brown, the areolet large, cubital and marginal cells 
closed. Abdomen densely black, highly polished. 

"Described from nine specimens, labelled No. 12.3, reared during 
July, 1SS6, by Mr. Albert Koebele, from Rhoditcs polila Ashm. and 
numerous other specimens labelled No. 38.39, reared at the Department 
during January and February, 188(5, from the same gall collected in 
Wyoming and Colorado," (W. H. Ashmead). 

I have not seen examples of this species. Doubtfully 

Periclistus obliquus Prov. 

Periclistus obliquus, Provancher, Addit. Hym. Quebec (1888), p. 397. 

"9 Long. .10 pec. D'un jaune saleavecune tache au metathorax, 
les jambes postericures, surtout a 1' extremite, et 1' abdomen en plus 
ou moins grande partie, noir. La face non striee. Les sillons parapsi- 
daux obliques, sc rapprochant en arriere. La radiale ouverte en avant, 
le radius atteignant le bord costal, 1' areole incomplete. Abdomen 
presque entierement noir; la tariere saillante, 1' ecaille ventrale temiinee 
en pointe fine. Los Angeles (Coquillett)." (L. Provancher). 

I have not seen examples of this species. 

Periclistus piceus n. sp. 
Female. Piceous black, mandibles, tcgulas, femora, tibia;, tarsi 
(except at tips), o\npositor and ON-ipositor sheath browTi or brownish. 
Head microscopically coriaceo-reticulate and pubescent, densely 
pubescent on face below antenna; and cheeks, antenuEB 1 2-segmented, 
filiform, 1st segment obconic, 2nd subglobose, stout, 3fd segment as 
long as 1 and 2 together, 3rd-6th subequal, following segments progres- 
siveh' shorter, except the last, which is more than twice as long as penul- 
timate. Pronotum and mesonotum faintly rugose, thickly clothed 
wdth appressed, whitish hairs, pronotum wide in the middle, parapsidal 
grooves on mesonotum incomplete, reaching less than half-way to 
anterior margin, median longitudinal lines and lines over base of wings 
inconspicuous, mesopleura smooth and shining, pubescent above and 
beneath, scutellum deeply rugose, pubescent, basal iovese small, cir- 
cular, shallow, with smooth bottom, contiguous, separated by a carina, 
metanotum pubescent. Abdomen smooth and shining, 2nd tergite 
covering the greater part of the abdomen, following tergites very narrow, 
posterior margin pubescent, ventral valv^e conspicuous, tuberculate. 

1911] CynipidcB of California 375 

sheath of ovipositor projecting upward obhquely to line of tergum. Legs 
pubescent. Wings hyaline, pubsecent, veins fuscous, distinct, radial 
cell short, closed, vein at base slightly arcuate, areolet distinct, cubitus 
not reaching basal vein. Length 2.5 mm. 

Male. Similar to female, but antennae 14-segmented, 3rd segment 
basally emargined, parapsidal grooves in many specimens apparently 
complete, pubescence more or less sparse. 

Bred from galls of Rhoditcs politus on wild rose (Rosa 
■calif ornica) . 

Habitat. Pt. Arena, Cal. (Miss Mabel Patterson.) 
Very similar to P. californicus, but apparently distinct. 


Diastrophus, T. Hartig, Zeils. f. Ent., vol. 2 (1840), p. 180, liU. 

Face radiately striate, antenna; in female 1.3-15 segmented, in male 
14-15 segmented, pronotum narrow in the middle, mcsonotum smooth 
and shining, bare, parapsidal grooves complete, distinct, scutellum with 
basal foveas. Radial cell open at the margin, claws bidentate, ventral 
valve scarcely as long as wide. Sexual. Galls on Rubus, Potentilla 
and Smllax. 

Diastrophus kincaidi Gillette. 

Diastrophus Idncaidii, Gillette, Can. Ent., vol. 25 (1893), p. 110. 

Diastrophus kincaidi, Kieffer, Bull. Soc. Metz, 2nd ser., vol. 10 (1902), p. 92. 

Female. Black, the antenna?, legs (except tips of tarsi), tcgulae 
and sheath of ovipositor brown or brownish. Head smooth and. shining 
on occiput, vertex and front to insertion of antennae, face radiately 
striate and pubescent, pubescence extending on to gense, antenna 
13-segmented, 1st and 2nd segments stout, 1st obconic, 2nd globose, 
3rd segment long, as long as 1 and 2 together, following segments progres- 
sively shorter to last, which is considerably longer than penultimate; all 
the segments more or less pubescent, distalljr from the middle infuscate. 
Pronotum wide in the middle, aciculate at the sides, pubescent, meso- 
notum smooth and shining, parapsidal grooves complete, rather widely 
separated at scutellum, median longitudinal lines and smooth lines over 
base of wings distinct, mesopleura transversely aciculate and shining, 
scutellum evenly rugose, basal fovea; oblique, oval, with smooth shining 
bottom, contiguous, separated by a median carina, metanotum rugose 
and bare. Abdomen smooth and shining, dorsally depressed, 2nd tergite 
reaching more than half-way to apex, 3rd tergite rather broad, oviposi- 
tor slightly protrtiding. Legs pubescent. Wings hyaline, pubescent, 
faintly iridescent, veins brownish, radial cell open at the margin, vein 
at base almost straight, areolet distinct, cubitus reaching basal vein. 
Length 3 mm. 

The male has 14-segmented antenncc, of which the 3rd segment is 
basally emargined. 

Gall. Large galls surrounding the stem of the thimbleberry {Rubus 
nutkanus), 25-60 mm. long and 12-25 mm. in diameter. Polythalamous. 

Habitat. Pt. Arena, Cal. (Miss Mabel Patterson.) Alameda 
County, California. (Beutenmiiller.) 

376 Annals Entomological Society of America [Vol. IV, 


Compsodryoxenus, Ashniead, Proc. U. S. Nat. Mus., vol. 19 (1896), p. 128. 

Head confluently punctate or faintly rugose, antermae filiform, 13-14 
segmented, thorax closely and confluently punctate or faintly rugose, 
parapsidal grooves shallow but distinct, pleura punctate, scuteUum 
rugose, abdomen compressed, ventral valve prominent, sharph- pointed, 
plowshare-shaped. Radial cell open, vein at base arcuate and sur- 
rounded by a brown cloud, margin of basal vein clouded and a brown 
spot before the break in the anal vein. Claws simple. Galls on Quercus. 

Compsodryoxenus bnuineus Ashm. 

Compsodryoxenus brunneus, Ashmead, Proc. U. S. Nat. Mus., vol. 19 (1896), 
p. 129. 

"Galls. The gall of this species was likewise confused in the collec- 
tion with a similar gall {Audricus chrysolepidis) occurring on Q. chrys- 
lepis in California, but I can distinguish two kinds of galls, although 
both bear the same number (2972). Both are very much alike exter- 
nally, but one is polythalamous, the other monothalamous, and I 
believe the latter is the one producing the present gall-fly. 

"Gall-fly. Female. Length, 2 to 2.6 mm. Head, antennas, 
thorax, and legs pale or light brown, the antennas towards tips dusky, 
pleura blackish, the abdomen black, polished, the posterior legs dusky 
or darker than the other. Wings hyaline, marked as in previous 

"The species is closely allied to the preceding, but it is smaller, 
paler colored, and has 13 joints in the antennas. 

"Type No. 30S5, U. S. N. M. 

"Specimens reared June 9, 18S3, (?1<S93). Under this number the 
record book contains the following; January 13, 1S93, Received 
today from Mr. H. W. Turner, of Martinez, California, a lot of elongate, 
oval t-nag galls, found on scrub oak; some of them were collected January 
3rd in Pine Canyon, Mt. Diabalo Contra Costa County, and some from 
apparently the same species of oak at Martinez ; placed galls from differ- 
ent locaHties into different bottles to breed. " (W. H. Ashmead). 

I have not seen examples of this species. 

14 RHODITES Hartig. 

Cynips (part.), Linne, Syst. Nat., ed. 10 (1758), p. 343, 553. 

Rhodites, Aylax (part.), T. Hartig, Zeits, f. Ent., vol. 2 (1840), p. 186, 194. 

Head large, broader than the thorax, antennas 14-15 segmented, 
pronotum narrow in the middle, parapsidal grooves complete or not 
reaching anterior margin, deep longitudinal groove beneath on meso- 
pleura, scutellum -with basal foveas or a ■wide transverse groove, abdomen 
smooth in female, strongly compressed apically, steniite of last segment 
plowshare-shaped, broad at the base and gradually narrowing to a point, 
abdomen in male small, compressed and rounded apically. Radial cell 
closed, short. Claws simple. Galls on Rosa. 

1911] Cynipida of California 377 

Rhodites bicolor (W. Harr.) 

Cynips bicolor, T. W. Harris, Treat. Ins. N. Engl. (18-42), p. 399. 
Rhodites bicolor, Osten-Sacken, Proc. Ent. Soc. Philad., vol. 2 (1S62), p. 
43, 48. 

Rhodites spinosellus, Cockerell, Entomol., vol. 23 (1890), p. 75. 

Female. Black, tips of mandibles, palpi, ocelli, legs (except coxje, 
trochanters and tips of tarsi), tegtila? and abdomen reddish brown. 
Head faintly punctate on occiput, vertex and front to insertion of 
antenna, face and genje coarsely punctate and clothed wdth silvery 
pubescence, antennas 14-segmented, 1st and 2nd segments stout, 2nd 
globose, 3rd segment longest, much longer than 4th and nearly twice as 
long as 1 and 2 together, succeeding segments to 9th progressively 
shorter, 9th and following seginents subequal, except the last, which 
is much longer than penultimate; all the segments rather thick. Prono- 
tum nan-ow in the middle, pimctate and pubescent, mesonotum elevated, 
rugoso-punctate, faintly pubescent, parapsidal grooves reaching more 
than half-way to anterior margin, median groove from posterior margin 
shorter, not extending beyond the middle, median longitudinal lines 
and smooth lines over base of wings distinct, mesopleiira rugose and 
faintly pubescent, smooth, bare patches in the middle and beneath, 
scutellum deeply rugose, basal fovea? wanting, sharp declivit\- on either 
side at base smooth and shining, metanotum rugose and pubescent. 
Abdomen compressed, smooth and shining, 2nd tergite reaching half-way 
to apex, 3rd and following tergites fairly wide, valves touched with 
black. Wings subhyaline, pubescent, iridescent, veins brownish, radial 
cell short, closed, vein at base angulatc, areolet distinct, cubitus reaching 
basal vein. Length 5 mm. 

Male. Similar to female but antenns and abdomen piceous black, 
legs more or less fuscous, wings hyaline, nen,^ures black. 

Gall. Spiny galls occurring in clusters on the wild rose {Rosa 
californica) ; yellowish brown, body spherical, the spines as long as or 
longer, sometimes shorter than the diameter of the body. Monothal- 

Habitat. Stevens Creek, above Cupertino, Cal. (R. W. 

Rhodites politus Ashm. 

Rhodites poHta, Ashmead, Bull. 1, Colo. Biol. Assoc. (1800), p. 14, 38. 
Rhodites politus, BeutenmuUer, Bull. Am. Mus. Nat. Hist., vol. 23 (1907), 
p. 644. 

Female. Black, the legs reddish bro\\Ti outwardly from base of 
femora. Head faintly punctate, face pubescent, antennee 14-seginented, 
filiform, 3rd segment longest, nearly twice as long as 4th or 1 and 2 
together, following segments progressively shorter, except the last, 
which is nearly twice penultimate. Pronotum narrow in the middle, 
closely pmictate and pubescent, mesonotum sparsely and shallowly 
punctate, each puncture bearing a short hair, parapsidal grooves com- 
plete, median longitudinal lines rather short, mesopleura finely rugoso- 
punctate, the disc bare and highly polished, scutellum rugoso-punctate 

378 Annals Entomological Society of America [Vol. IV, 

much longer than wide. Abdomen smooth and shining, compressed 
beneath, ventral valve projecting and acutely pointed. Wings hyaline, 
pubescent, veins brownish black, radial cell short, closed, with a brown- 
ish cloud, vein at base angulate, areolet distinct, cubitus reaching 
basal vein. Length 3 mm. 

Gall. Small globular galls with weak spines, on leaves of wild rose 
{Rosa calif ornica), often in tangled clusters. 

Habitat. Pt. Arena, Cal. (Miss Mabel Patterson.) Los 
Angeles. (Beutenmiiller.) 


Lytorhodites, Kieffer, Bull. Soc. Metz, ser. 2, vol. 10 (1902), p. 96. 

Differs from RkoJiles only in the following particulars; scutellum 
without fovejE, radial cell more or less open at the margin, abdomen 
usually faintly reticulate. Galls on Rosa. 

Lytorhodites arefactus (Gillette). 
Rhodites arefactus, Gillette. Can. Ent., vol. 26 (.1894), p. 1.57. 
Lytorhodites arefactus, Kieffer, Bull. Soc. Metz., ser. 2, vol. 10 (1902), p. 97. 
Rhodites similis, Beutenmuller, Bull. .'\m. Mus. Nat. Hist., vol. 2.3 (1907) 
p. 640. 

"The galls arc dense, corky enlargements of small shoots, tisually 
close to the stem from which the shoot arises, and the shoot is usually 
dead beyond the gall when the latter is mature. The galls are irregular 
in shape, vary from one-half to seven-eighths of an inch in diameter, 
and are polythalamous. The surface is of a rusty color, is finely 
wrinkled, and reminds me of dried fruit. The surface appears dry and 
hard, but it is easily dented with the finger-nail and is always free 
from spines. 

"Described from eighteen galls collected in March in the vicinity 
of Fort Collins, Colorado. Galls brought into the laboratory March 
7th began giving flies March 23rd. 

"Gall-flies. Females. General color cinnamon-rufous; head en- 
tirely rtifous, except a blackish area between either compound eye and 
the mouth ; under a power of 60 diameters the lower face appears rather 
coarsely wrinkled, the wrinkles converging towards the mouth, the upper 
face, vertex and occiput very finely rugose; the face sparsely set with a 
short gray pubescence; antennas .short, the first three joints, and some- 
times the base of the third, rtifous, the remaining joints black; number 
of joints, 14. Thorax rtifous above, with a black suture separating 
the mesothorax and scutellum, parapsidal grooves entire, broad, moder- 
ately deep, well separated at the scutellum, and with numerous ele^^ated 
lines crossing them; median grooves distinct and extending well forward. 
The surface of the thorax is finely rugose, and, in a proper light, shows 
numerous punctures, each puncture bearing a short yellowish hair. 
Scutellum coarsely wrinkled near the margin and less coar.sely wrinkled 
on the central portion, which is considerably elevated, transverse 
groove at base, color rufous. Mesopleura, except spot just beneath 
the ■n'ings, rufous, sutures, metathora.x and sternimi black or blackish; 


CynipidcE of California 


entire pleura rugose. Abdomen rufous, with venter and posterior 
half of dorsum blackish, all abdominal segments covered with a micro- 
scopic network of impressed lines, most prominent on the terminal 
segments. Wings but little smoky, radial area not at all closed along 
the costal margin, areolet distinct and rather small. Feet, including 
the coxae, entirely rufous, the claws onlv being black. Length froin 
3K to iyi mm. 

"Described from twenty-one specimens bred from the galls. 

"Males. Three to three and one-half mm. in length, black, feet 
more reddish in color than in the females, bases of the coxae black, 
antennas black throughout; otherwise like the females. 

"Described from forty-two bred specimens. 

"There is one male among those reared that has the rufous marking 
of the female on head, antennae and thorax. 

"This species resembles very closely Rhodites muUispinosa Gill., 
but the galls are very different." (C. P. Gillette). 

I have not seen examples of this species. 


agrifoliae (Synergus) 

agrifolia (Andricus) {Callirhytis) . . . 


apicalis (Callirhytis) 

apicalis (Andricus) 

arefactus (Lytorhodites) . 

aref actus (Rhodites) 

baiceri (Callirhytis) 

batatus (Neuroterus) 

bicolor (Rhodites) 

bicolor (Cynips) 


brevicomis (Synergvis) 

brunneus (Andricus) 

brunneus (Compsodryoxenus) 

calif ornica (Biorhiza) 

californica (Philotiix) 

calif ornicus (Periclistus). . . 

californicus (Andricus) (Callirhytis) 


canescens (Cynips) 

canescens (Holcaspis) 

canescens (Disholcaspis) 


chrysolepidicola (Callirhytis) 

chrysolepidis (Disholcaspis) 

ckrysolcpidis (Holcaspis) 

chrysolepidis (Andricus) 

clarimontis (Callirhytis) 

clavula (Diplolepis) 

clavula (Dryophanta) 


congregatus (Andricus).. . 

364 coquilletti (Trichoteras) 3-11 

356 corallina (Cynips) 343 

346 corallina (Disholcaspis) 343 

354 corallina (Holcaspis) 343 

.354 crystallinus (Andricus) 348 

378 Cj-nips ,. .342 

378 dasydactyli (Andricus) . . 349 

360 Diastrophus , 375 

334 dimorphus (Synergus) . - - 366 

377 Diplolepis 336 

377 discus (Diplolepis) 336 

333 discus (Dryophanta) 336 

365 Disholcaspis 339 

353 dorsalis (Ceroptres) 373 

376 douglasi (Diplolepis) 338 

334 douglasi (Dryophanta) 338 

334 douglasi (Holcaspis) 338 

.374 Drvophanla 336 

346 dubiosa (Diplolepis) 339 

354 dubiosus (Synergus) 372 

342 echina (Diplolepis) 337 

342 echina (Drvophanla) 337 

342 echinus (Cynips) 337 

372 eldoradensis (Disholcaspis) .340 

354 eldoradensis (Holcaspis) 340 

341 eriophora (Callirhytis) 359 

341 flavus (Synergus) 365 

346 flocci (Cynips) (Andricus) 352 

359 flocci (Andricus) 3.52 

3.37 guadaloupensis (Callirhytis) 363 

337 hclda? (Cynips) 345 

376 Holcaspis 339 

.347 kelloggi (Cynips) 345 


Annals Entomological Society of America [Vol. IV, 

kingi (Andricus) 350 

kincaidi ( Diastrophus) . ' 375 

lana (Cynips) 352 

lana (Atidrkus) 352 

lasia (Callirhytis) 356 

Lytorhodites 378 

maculatus (Synergus) 371 

maculipennis (Cynips). 344 

maculipennis (Callirhytis) 358 

maculipetinis (Holcaspis) 344 

maculipennis (Disholcaspis) 344 

multiplicatus (Synergvis) 370 

multipunctata (Cynips). , . 343 

mullipunclata (Dryophanta) 343 

Neuroterus 334 

niger (Ceroptres) 373 

niger (Synergus) 369 

nigra (Callirhytis) 362 

obliquus (Periclistus) 374 

ochreus (Synergus) 368 

oneralus (Cynips) 366 

oneratus oneratus (Synergus) 366 

oneralus (S^'nergus) 366 

pacificus (Andricus) 348 

parmula (Andricus) 350 

pattersonas (Andricus) 352 

Periclistus 373 

Philonix 333 

piceus (Periclistus) 374 

politus (Rhodites) 377 

polythyra (Callirhylis) 357 

pomiformis (Ceroptres) 372 

pomiformis (Andricus) 355 

pomiformis (Callirhylis) 355 

punctatus (Sjmerg\is) . 367 

quercus-agrifolis (Callirhytis) 356 

quercus agrifolia (Cynips) 356 

quercus batalus (Cynips) 334 

quercus-batatus (Neuroterus) 334 

quercus californica ( Cynips) 346 

quercus-califomicus (Andricus). . . . 346 

quercus flocci (Cynips) 352 

qucrcus-flocci (Andricus) , . 352 

quercus lana (Cynips) 352 

quercus pomiformis (Cynips) 355 

quercus-pomiformis (Callirhytis). . 355 

quercus suttoni (Cynips) 357 

quercus suttoni (Callirhytis) . . . 357 

Rhodites . 376 

rossi (Callirhytis) 361 

sallatorius (Cynips) . 335 

saltatorius (Neuroterus) 335 

sanctae-clarae (Callirhytis) 363 

similis (Rhodites) 378 

speciosa (Dryophanta) 337 

spinoscUus (Rhodites) 377 

splendidus (Synergus) 369 

suttoni (Andricus) (Callirhylis) .. 357 

Synergus . 3(>4 

Trichoteras , 341 

truckeensis (Disholcaspis) 340 

truckeensis (Holcaspis) 340 

vaceiniifolice (Callirhytis) 357 

varicolor (SjTiergus) 371 

wiltzEe (Andricus) 353 

wisliceni (Andricus) 337 

Annals E. S. A. 

Vol. IV, Plate XXIII. 




(Coleoptera, Family Curculionidae) 


By E. G. Titus, Sc. D. 


Introduction 383 

Acknowledgments '. 384 

History o£ the genera and group 386 

Characters of tribe Hyperini 389 

Life-history of the group 391 

Food-plants of the Hyperini in America 393 

Habits, injuries and control 393 

Genera Hypera and Phytonomus, characters of 394 

Table of species 395 

Hypera punctata 396 

Phytonomus cximius 412 

Phytonomus quadricollis 414 

Phytonomus comptus 415 

Phytonomus diversipunctatus 421 

Phytonomus seriatus 426 

Phytonomus trivittatus 428 

Phytonomus maritimus 432 

Phytonomus castor 433 

Phytonomus pubicollis 434 

Phytonomus meles 435 

Phytonomus nigrirostris 442 

Phytonomus posticus 452 


The genera Hypera and Phytonomus belong to the tribe 
Hyperini, a member of the subfamily Curculioninae of the 
Coleopterous suborder Rhyncophora. 

Both genera are well distributed over the northern hemi- 
sphere, being especially abundant in Europe, the last catalog 
of Heyden, Reitter and Weise listing in Phytonomus from 
"Europae caucasi et Armeniae Rossicae" 64 species, 3 varieties 
and 21 aberrations. 

In America Leconte in 1876 listed 9 species, three of which 
are European. Since that time, we know of the introduction 
of two more European species. The present paper includes 
13 species. One Phytonomus has been described from Mexico 
and in South and Central America are a number of species of 
Phelypera, a very closely related genus. 

In the present paper will be treated only those species 
known to occur in America, north of Mexico, of the genera 

* Revised from a thesis submitted as a partial requirement for the degree of 
Doctor of Science at Harvard University, May 1, 1911. 

Contributions of the Entomological Laboratory, Bussey Institution, 
Harvard University, No. 39. 



Annals Entomological Society of America [Vol. IV, 

Hypera and Phytonomus. Specimens have been seen of all the 
species reported from this region, and the types of six species 
have been examined. 

In connection with the study of the American species the 
author has studied 45 of the European species, in some cases 
including .a large number of specimens. Over 500 American 
specimens have been examined, exclusive of several thousand 
specimens of P. posticus. 


It is with pleasure that I here acknowledge the generous 
loans and gifts of material and the receipt of many records 
from the various sources here given. 

AIap 1. Distribution of Hypera and Phytonomus. 

From the personal collections of C. A. Frost, Framingham, 
Mass.; F. A. SherrifY, Melrose Highlands, Mass.; Frederick 
Blanchard, Tyngsboro, Mass. ; C. T. Brues, Bussey Institu- 
tion, Harvard University; Charles Schaefler, Brooklyn, N. Y. ; 
R. P. Dow, New York, N. Y.; E. A. Bischofif, Irvington, N. J.; 
Henry Wenzel, Philadelphia, Pa.; W. S. Blatchley, Indianapo- 
lis, Ind.; J. D. Evans, Trenton, Ont.; A. B. Wolcott, Chicago, 
111. ; Prof. H. F. Wickham, Iowa City, la. ; R. L. Webster, Ames, 
Iowa; Warren Knaus, McPherson, Kan.; Franklin Sherman, 
Jr., Raleigh, N. C; Norman Criddle, Aweme, Man.; Trevor 

1911] Ilypera and Phytonomiis in America 385 

Kincaid, Seattle, Wash.; G. I. Reeves, Pullman, Wash.; E. C. 
Van Dyke, vSan Francisco, Cal. ; F. E. Blaisdell, San Fran- 
cisco, Cal.; Henry C. Fall, Pasadena, Cal.; and C. N. Ainslie, 
Salt Lake City, Utah. 

Determined European material of Phytonomits miiriniis, P. 
variabilis and P. viciae, has been received from Dr. Edmund 
Reitter of Paskau, and Prof. Victor Ferrant of Luxemburg, 
and numerous specimens representing a number of European 
species from Baron von Rothkirch, Lubben, Germany. 

From the following institutions I have had material and 
records: U. S. N. Museum through Dr. L. O. Howard and Mr. 
E. A. Schwarz; Am. Mus. Nat. History, Mr. Frank Lutz; 
Department of Agriculture, Dominion of Canada, through 
Dr. Gordon Hewitt; Brooklyn Academy Arts and Sciences, 
Charles Schaeffer; American Entom. Society and Philadelphia 
Academy of Sciences, Dr. Henry Skinner; State Entomol- 
ogist's office of New York, Dr. E. P. Felt; Cornell University, 
Dr. Alex. MacGillivray; Illinois University (Bolter collection) 
and 111. St. Lab. Nat. History, Dr. S. A. Forbes; Field Colum- 
bian Museum, W. J. Gerhard; Colo. Agric. College, Prof. 
C. P. Gillette; Conn. Agr. Exp. Station, Prof. W. E. Britton; 
N. J. Agric. College, Dr. J. B. Smith; Boston Soc. Nat. Hist., 
Mr. C. W. Johnson, and last but by no means the least I have 
had the advantage of studying the collections in the Museum 
of Comp. Zoology at Cambridge, Mass., and the kind and ever- 
ready aid of Mr. Samuel Henshaw, Curator, in searching the 
literature and examining specimens. 

To Dr. L. O. Howard I am indebted for the translation of a 
paper by Dr. Martelli that was published in March of this year. 

To Mr. C. T. Brues and Mr. F. W. Chapman, for aid in 
securing the photographs of many of the species here 

To Prof. F. M. Webster, of the Bureau of Entomology, 
Washington, D. C, and his assistants, who have aided in the 
alfalfa weevil work in Utah. 

To my associates in Utah, and especially to Dr. E. D. Ball, 
Director of the Experiment Station, for having made possible 
the opportunity to study the life history, and to my assistant, 
Mr. V. A. Sadler, for his efficient aid in the field work on 
Phytonomiis posticus. 

386 Annals Entomological Society of America [Vol. IV, 

Finally, I am greatly indebted to Dr. W. M. Wheeler, of 
Bussey Institution, Harvard University, under whom much of 
the systematic work has been developed, for his sincere kind- 
ness, his encouragement and advice. 


In 1817 Germar in Germar & Zincker's magazine published 
a short article calling attention to the fact that he had for a 
long time been studying the genus Curculio, and that he had 
found good characters in the large complex of species upon 
which to erect new genera. At that time he published the 
names of these proposed genera, each with one or more included 
species, promising later to give the descriptions. 

In 1821, in the same magazine, he published descriptions of 
several of the genera noted in 1817. Among these was the 
genus Hypera, which he divided into two groups, containing 
altogether 14 species. In the former paper he gives no char- 
acters whatsoever to distinguish the different groups, merely 
mentioning some of the work he had done and giving the list. 
The genus dates from this latter paper (1821). W'-'^ 

In 1826 Curtis in his illustration of British Insects figured 
on plate 116, dated May 1, 1826, Ilypera Jasciculala, and stated 
in the appended description that the type of the genus Hypera 
was Curculio punctatiis. He included in his list a number of 
other species which he had examined. //. punctatiis was one 
of the species included by Germar in the original description of 
the genus and hence will stand as the type. 

In the same year (1826) Schonherr published his work 
" Curculionidum dispositio methodica," in which in pt. iv, p. 175, 
"he erects the genus Phytonomus, dividing it into two groups, 
nearly identical with those of Hypera given by Germar. He 
makes the type of the genus and of his first group Hypera 
polygon i L. Hyp. punctata was included in his second group. 
He gave with each group a number of species which he consid- 
ered as belonging to that complex. He undoubtedly intended 
to make Hypera a complete synonym, but since the type of 
Hypera had already been fixed, both genera should stand. At 
later dates both Gyllenhal and Germar accepted the genus 
Phytonomus as including all the species under the two groups, 
upon what grounds it is impossible to state. Giebel cites the 
ispecies in the collection at "Univ. Halle-Wittenberg" under 

1911] Hypera and Phytonomus in America 387 

the name of Hypera. It is apparent that these incktde Germar's 
material and would indicate that he had not changed the name 
in his own collection. 

Why later writers (Jekel, Lacordaire and especially Fowler) 
should attempt to fix other types for the genus Hypera I do not 
know, unless I have overlooked papers to which they had 
access. There is no clue to such literature in their articles. 

Capiomont in 1867-8 in his "Revision des Hyperides," 
accepts this group as outlined by Lacordaire in the Genera des 
Coleopteres, tome vi, p. 395. Capiomont creates several new 
genera in the group and separates Phj^tonomus from Hypera, 
but not on the lines given by Curtis. The paper is, however, 
very valuable, as being the first thorough treatment of the 
group after Schonherr. 

Kirsch and Kraatz, each publishing in 1871, contribute 
nothing new to the separation of the groups included, neither 
does Seidlitz in his Fauna Transylvanica in 1891. 

Petri in 1901 in his admirable monograph of the tribe 
Hyperini closely followed Cai^iomont's work. He also gives a 
very good list of the synonomy of the species. 

The larger European catalogues before 1901 usually treated 
the genus Hypera with Phytonomus as a synonym or a sub- 
genus. Weise in the Heyden, Reitter and Weise Catalog of 
1906 has followed Petri except in the synonomy of the species, 
where he recognizes "aberrations" for most of those forms 
previously called varieties. This is certainly a step in the 
right direction, since in the species I have studied these so-called 
aberrations appear to be nothing more than forms due to one 
of several causes and likely to appear in any generation of the 
species. The term evidently should cover all such cases as. 
immature specimens, color changes due to temperature or food 
conditions, size forms, and specimens which have lost all or 
part of their pubescence, especially the scales. Most of these 
should never have been described, as is evident from the efforts. 
of both Capiomont and Petri to separate such forms from the- 
typical species as they conceived it. 

The literature of the group is of considerable size as may 
be judged from the bibliography of the species here treated. 
Without doubt references have been omitted that should have 
been given, but I trust that no serious omissions occur. The 
effect of this large number of references has been to make the 

388 Annals Entomological Society of America [Vol. IV, 

synonomy' of the species very difficult. It is apparent that the 
name of Phyto)iomns melcs is in doubt, but I do not care to 
change it without knowing more regarding the species trivialis 
Herbst and rocseli Gmelin, both of which were described 
previous to Fabricius' description of meles. The species 
major Herbst which had been assigned here as a synonym is 
according to Schonherr a Cleonus. Schonherr in his monu- 
mental work on the Curculionids in 1834 and 1842 gives no 
further aid on this synonomy. 

Regarding the Stephens species the papers of Walton have 
been followed even where they differ from later authors since 
it is believed that they more nearly represent the true synonomy. 
Walton was in correspondence with Germar and Gyllenhal and 
with other continental European entomologists, and exchanged 
specimens with them. Where there was a further question, 
the papers of Capiomont and Petri have been followed if 

I have not attempted to place the American forms in the 
subgenera given by Capiomont believing that these need so 
much revision as to names and species included that it is well 
to let them alone. A more thorough study of the life-history 
of the various Eurasian species will without doubt introduce 
more synonomy if the other species of that region are as 
variable as those introduced into America. 

The most constant characters are the scale structure, form 
and shape of thorax, shape and size of beak, and the genitalia. 
Petri has well pointed out the extreme differences in the stem 
of the male genitalia, the "forceps" of Petri. 

All the species described from America north of Mexico 
have been identified, and I feel compelled to make one of the 
Leconte species, Phy. setigerus, a synonym of trivittatiis of Say, 
this latter not having been previously recognized since Say's 
description. One new species, Phy. maritimits is described. 
Hypera ocellata, 1902: (Biol. Cent. Am. Coleop. v. 4, pt. 4, p. 3) 
was described from "Omilteme, Guerrero, 8,000 feet, Mexico 
(H. H. Smith)." From the description and figure, it apparently 
belongs near Phytonomus eximius. 

The term Phytonomini of Leconte must give way to 
Hyperini, the genus Hypera being erected prior to Phytonomus. 

1911] Ilypera and Phylonomus in America 389 


1863: Lacordaire; Gen. des Coleopteres, tome vi, p. 395 (Hyperides). 

1867: Capiomont: Revision de la Hyperides (Ann. Soc. Ent. Fr. pp. 417-560, 

pi. 11-12). 
1868: Capiomont: Rev. de la Hyperides (con.) (Ann. Soc. Ent. Fr. pp. 7-4-284, 

pi. 1-4). 
1871: Kirsch: Zur Kenntnis der deutschen Hvperiden (Berl. Entom. Zeits., 

pp. 173-191). 
1901: Petri: Monogr. des Coleop. — Tribus Hvperini, pp. 210, figs. 58, pi. 3, also as: 

Bestim.— Tab. Coleop. Hft. 44, pp. i-42. 

In this group the body is more or less oval, the thorax 
never exceptionally long, the beak never extremely long and 
slender; thorax and elytra more or less covered with scales and 
with hairs that may be simple, emarginate or thickened. 

The head is small and round, with the beak or rostrum well 
developed, often with a carina or keel on the upper surface; 
antennae set in a groove on the side of the beak, the groove 
usually slopes downward toward the lower side of the eyes; 
antennae (PI. XXIV, fig. 17), composed of twelve joints, a long 
scape, seven funicle joints, the first two of which are longer 
than any of the others, and a four-jointed club, the antennae 
rarely reach to the middle of the prothorax; eyes oval, round or 
elongate-oval, often narrowed below, rather large and close 
together in front; mouth parts at the apex of the beak as usual, 
labrum wanting, mandibles, (PI. XXIV, fig. 3, 4, 15), often with 
fine punctures, broad, stout, more or less pincer-shaped, with 
teeth, maxilla (PI. XXIV, fig. 1, 16) broad, with short, conical 
four-jointed palpi which are rigid and taper more or less to a 
point; lacinia provided with stout teeth and rather long hairs, 
apparently always with short spines on the inner surface; 
subntentum nearly rectangular, emarginate; mentum short and 
broad, labial palpi (PI. XXIV, fig. 2) three-jointed, rigid, 

Prothorax more or less rounded above as seen from the side, 
sides usually somewhat swollen, anterior and posterior margins 
rarely as wide as the middle; oval, transverse-oval or elongate; 
always with a short process below between the front coxae. 

Scutellum always minute. 

Elytra differing greatly in form, sometimes at least three 
times as long as the thorax, in other species less than twice as 
long, elongate; oval, broad or obovate; from the side usually 
rather flat at the base, often rising for a short distance, and 
then may be abruptly declivous or rounded to the apex; 10 
striae and 11 interspaces including the sutural and side spaces. 

390 Annals Entomological Society of America [Vol. IV, 

Venter as in other Curculionidae, front coxae almost con- 
tiguous, middle coxae separated by the more or less elevated 
process of the mesosternum and the shorter process of the 
metasternum; hind coxae usually rather widely separated, the 
intercoxal process of the third abdominal (first visible) seg- 
ment being broad, but in all species examined ends in a point 
which is sometimes concealed beneath the metasternum; side 
pieces of the mesosternum diagonally divided; side pieces of the 
metasternum dilated in front usually rather narrowly, the 
outer angle causing a sinuousity in the edge of the elytron; 
ventral abdominal segments unequal, first and second con- 
cealed as usual, the fifth and sixth shortest, usually the seventh 
or fourth next, the third longest, or in some the seventh the 
longest; sutures straight or nearly so. 

Last dorsal abdominal segment in the male with an extra 
piece, which appears as another segment and is (in the species 
examined) covered with peculiar many branched hairs or scales, 
pygidium not exposed. 

Legs clothed with hairs or scales, usually the femora and 
coxae with scales only, sometimes these only in front ; the apex 
of each of the tibia possesses a ring or crown of spines of varying 
length; articular surface of the hind tibiae distinctly terminal, 
sometimes with a projection on the inner side; tarsi dilated, 
third joint strongly bilobed, elongate, with a setose pad 
beneath; claws long, simple, free. 

The description of the stages relate only to the following 
species: Hypera punctata, Phytonomus posticus, P. nigrirostris, 
P. meles, and P. comptus. The characters seem however, to 
be common to the species named, where the stages are known. 

Egg: (not known in comptus) : more or less oval, white, 
yellow or some shade of yellow, reticulated with hexagonal 

Larva: In the younger stages (not known in comptus) 
slender, widest in the middle, head dark, dorsal surface set 
with dark or black tubercles upon which are inserted hairs, 
which are usually clavate at the tips, except on the anal seg- 
ments where they are longer and simple ; beneath the thorax the 
surface is projected into lobes, sometimes each lobe of each of 
the three pairs is bilobed and set with bristles; abdominal 
segments beneath with smaller lobes; sides with two swollen 
areas on each segment, the one on which the spiracles are placed 

1911] Ilypera and Phytonomiis in America 391 

has one or more tubercles set with hairs, anal segment of three 
lobes, two side and one dorsal; a dorsal abdominal median 
paler line is present, this may extend onto the thorax. 

Later stages: Head dark, glabrous with very fine transverse 
lines, antennae minute, two-jointed, situated near the anterior 
border; labrum emarginate, with a row of hairs near the edge; 
mandibles stout, toothed, usually dark; two small ocelli on 
each side of the head with a long hair between them; palpi 
two-jointed, a long hair or spine below the first joint. Seg- 
ments of the body dorsally of two distinct parts, (PL XXIV, figs. 
23-32), the smaller anterior part always with one pair of 
tubercles, a tubercle each side of the dorsal line; the posterior 
part larger, broader and extending almost to the spiracles, 
containing on the dorsum, at least, one row of tubercles, some 
of the thoracic and last abdominal segments may have more 
rows; spiracles black, nine in number situated a little above the 
middle and well forward on the side of the segments which 
possess them, below them one or two tubercles, the spiracles 
and these tubercles are on the first set of swellings or enlarge- 
ments; the enlargement below the first is usually small, the 
third is on the venter and contains the leg-like tubercles. 

Cocoon: All the species noted above spin reticulate cocoons, 
usually oval or globular, varying considerably in the size and 
shape of the openings. 

Pupa: Rather short and wide, all the appendages very evi- 
dent, wing-pads rather long, thorax broad, the abdominal seg- 
ments with transverse rows of setae; the thorax with hairs, 
those on the prothorax regularly twenty in number, a row of 
five pairs curving around the anterior margin on each side, the 
fifth of which is sometimes set far back; and a curved row of 
five pairs beginning near the center and passing backward to 
the posterior outer angle. The arrangement of these hairs 
appears to be constant in each species examined (pupa of 
meles not seen). 

Life-history: Eggs laid, except with Hyp. punctata, in the 
spring on the food plants or inserted into some part of the 
plant, such as leaf, leaf-sheath, petiole, stem, flower-heads or 
buds. The habit of comptus is not known, but from the time 
the larvae appear it is probable the eggs are laid in the spring, 
the same holds for P. eximius. 

392 Annals Entomological Society of America [Vol. IV, 

Larvae upon hatching, generally remain concealed for some 
time feeding in a protected place, usually not feeding in the 
open except at night or when very numerous, when they pass 
out onto the leaves even in broad daylight. Some feed in 
flower-heads {meles and nigrirostris, prob. also eximius and 
comptus) ; others in the leaf -buds {posticus) , but all when 
numerous will defoliate their food plant. 

The larva when full grown spins a cocoon that may be 
placed among the flowerets (nigrirostris) , on the upper surface 
of the leaves (comptus); on or near the ground, in leaves or 
other debris (posticus) or in the ground (H. punctata). Even 
in the same species there is some diversity of habit. 

In those species where the cocoon spinning has been watched 
the process is as follows (Folsom, Titus for Hyp. punctata, 
Titus, Ainslie, Sadler for P. posticus, Titus for P. nigrirostris). 

In Ilypera punctata the larva buries itself in a small oval 
cell in the ground, slightly under the surface; this cell it smooths 
with its head and by turning around and around with its body 
in- the characteristic curved position; the other species do not 
form cocoons in the ground. The spinning in the species 
observed is done with the mouth. The first hairs are placed as 
a round network on the surface where the larva is lying, then 
lying on its back it reaches with the head to one side slowly 
spinning the thread upward. The thread hardens and is thus 
sometimes carried over to the other side making a framework 
upon which to attach other threads. More often the threads 
are laid down along the first network and gradually built up on 
each side, the larva often puts its mouth or parts of its mouth 
through the coarser network and fastens a thread outside. The 
meshes are gradually reduced, in size by placing other threads 
in both directions inside the first rows, this is especially true 
with comptus and punctata. 

Every few seconds, or at least every half minute the larva 
reaches back to the anus and apparently from some gland 
secures a fresh supply of silk, the operation of securing this silk 
can be better described as sucking than "nibbling" though it 
partakes of the character of both. It may be that this is a 
secretion from the malphigian glands as found by Silvestri to 
occur in Lebia. Pupation occurs from one to three days after 
the cocoon is completed. 

1911] Hypera and Phytonomus in America 393 

When the adult beetle appears it rests in the cocoon until 
the wing-covers are somewhat hardened and then eats its way 
out. From the descriptions given apparently some species 
devour the entire cocoon, this has been noticed but rarely with 
posticus and has not been reported for punctata. 

The beetles usually feed by night and rest concealed in the 
daytime beneath rubbish or leaves or even in cracks in the 
ground. The smaller specimens often lie in the leaves or 
opening leaf-buds. The beetles cause considerable injury by 
their feeding habits at this time, gnawing the parenchyma 
from the stems and feeding upon the leaves. 

The introduced European species, and probably all the 
ispecies, hibernate as adults. 

The group has in common with some other Curculionidae 
the habit of distributing themselves by flying at some stated 
period, in Phytonomus it appears at least in three species 
{nigrirostris, posticus, meles) to be in the spring. P. posticus 
has two flights, the second occurring in the summer, Hypera 
punctata has at least late summer or fall flight. 

Food-plants: Kleine has published (1910j the food plants of 
the European species of Phytonomus so far as known. They 
include plants in many different groups, but especially among 
the legumes (Fabaceae) and buckwheat (Polygonaceae) families. 

The native American species whose food-plants are known 
are comptus on Polygotium; eximus and quadricollis on Rumex; 
trivittatus (setigerus Lee.) on Lathyrus, and maritimus on 

The introduced species • are primarily leguminous feeders, 
attacking especially clovers and alfalfa; probably they will feed 
upon any species of Trifolium, Medtcago or Melilotus. They will 
.also attack the Astragalus group and the vetches. Hyp. punctata 
will live upon beans in both larval and adult stages. Other food- 
plants reported for them, such as golden-rod, potatoes, timothy, 
wheat and cabbage, are doubtless more or less accidental. 

Along the Atlantic Coast the introduced species are but 
occasionally noticed as injurious to the crops, but as they move 
westward across the Alleghany Mountains, their injuries 
increase. It is probable that when Hypera punctata from the 
East and Phytonomus posticus from the West meet on the 
western plains, we will hear much more regarding their injurious 
feeding habits. It is certain that the alfalfa weevil (P. posticus) 

394 Annals Entomological Society of America [Vol. IV, 

is a most serious pest in the parts of Utah where it is at present 
common and doubtless will be so in any of the western alfalfa 
regions. Railroads lead in all directions and it is only a matter 
of time until this species has reached the other alfalfa growing 
localities. (Map 11.) 

In the dry regions, where there is little rainfall during the 
hot summers and very little humidity in the atmosphere, it is 
very doubtful if the fungus diseases will work. In the coast 
regions the fungus is undoubtedly the one enemy that keeps 
the species there present in check. 

Cultural methods, the introduction of better methods of 
farming, rotation of crops, use of gathering machines, careful 
stamping out of incipient colonies and the hope of parasites 
from Europe are all factors leading toward the control of the 
species of this group in the more arid climates. 

Plates XXXIII and XXXIV illustrate something of the 
problem from the standpoint of the western farmer and show 
what is being done to aid in cultural lines. Plate XXXIII is 
adapted from Bui. 110 of the Utah Agr. Exp. Sta., which gives 
an account of the work accomplished against P. posticus up to 
July 1, 1910. 


While these two genera are closely related, there are unmis- 
takable characters that readily separate them. Ilypera has 
the beak short, blunt and thick; scarcely one-half longer than 
the remainder of the head; mandibles never emarginate; 
elytra much wider than the thorax; usually wider or as wide as 
the distance from base to point where the elytra curve down- 
ward; humeri very prominent, convex; alternate interspaces 
beginning with the sutural one strongly elevated and wider 
than the others; intercoxal process broad, stem of male 
genitalia (PI. XXIV, fig. 14), fully as wide as long. 

Type: Hypera punctata Fab. 

In Phytonomus the body is never stout, broad and thick, 
beak never short and blunt; intercoxal process somewhat 
narrowed at tip; mandibles always more or less emarginate; 
elytra as wide or a little wider than the thorax; stem of male 
genitalia (PI. XXIV, figs. 5-13), much longer than wide, 
generally two or three times longer. 

Type: Phytonomus arator L. (polygoni L.) 

1911] Hypera and PJiytonomus in America 395 

Hypera lays eggs in the fall, some larvae hatch then, others 
the next spring. Cocoon may be formed in the ground, and the 
meshes are very much closer than in any Phytonomus cocoon 

Phytonomus lays eggs in the spring (so far as known) ; the 
cocoon apparently never formed beneath the surface of the earth. 


Beak stout, never longer than prothorax. 

Large robust species, beak shorter than prothorax; hairs of prothorax and 
elytra long and slender; scales striate, narrowed toward tip, cmarginate 

without processes, concave, rounded at base Hypera punctata 

Elongate, rather stout, sides of elytra almost parallel, thorax longer than 
broad, beak scarcely as long as prothorax; setae on prothorax thick, 
numerous, scales sparse, parallel-sided, deeply emarginate. 

Phytonomus diversipunctatus 
Beak more slender, always longer than prothorax. 

Front between the eyes narrower than eye at widest part 1 

Front between the eyes always distinctly wider than eye at widest part, 
usually slightly concave 8 

1. Scales not at all cleft or emarginate 2 

Scales more or less emarginate 4 

Scales cleft 6 

2. Body not elongate, flattened, sides of elytra never parallel; scales trun- 

cate, concave, widest at middle, striate Phytonomus eximius 

Body elongate, flattened, sides of elytra parallel 3 

.3. Thorax deeply punctured, polished; scales concave, truncate, widened at 
tip, finely striate; setae thickened at tip, more numerous on posterior 

part of elytra Phytonomus quadricollis 

Thorax not polished, punctures shallow, indistinct, glabnais, more or less 
confluent; scales parallel-sided or narrowed at tip. thick, iiulistinctly 
striate, no hairs on dorsum Phytonomus comptus 

4. Scales finely striate, deeply emarginate, sides curved; hairs thick at base 

and near tip abruptly narrowed to a point; prothorax as wide as long; 

punctures of elytral striae with minute setae . Phytonomus trivittatus 
Scales deeply emarginate; hairs on prothorax thick, sides parallel, tip 

notched; prothorax longer than wide, setae in elytral striae short, 

thick and white Phytonomus maritimus 

Scales deeply emarginate, processes and elytral scales as long as body of 

scale; beak scarcely longer than prothorax, species small, sto'.it. 

Phytonomus pubicollis 

5. Scales cleft nearly or quite to base 6 

Scales not nearly cleft to base 7 

6. Prothorax much wider than long, sides prominently rounded. 

Phytonomus meles 

Prothorax not wider than long, species rather narrow, elongate; hairs on 

dorsum long, fine, pointed Phytonomus nigrirostris 

7. Prothorax almost as wide as long; hairs on dorsum, especially on posterior 

part of elytra, semi-decumbent, long and pointed. .Phytonomus posticus 
S. Scales of elytra cleft to or almost to base; prothorax with numerous 
emarginate hairs mixed with sparse cleft scales, front distinctly eon- 
cave, beak much longer than prothorax Phytonomus seriatus 

Scales of thorax and elytra all deeply cleft, some scales on head are cleft; 
prothorax with sparse, thickened, blunt hairs, beak scarcely longer 
than prothorax Phytonomus castor 

396 Annals Entomological Society of America [Vol. IV^ 

Hypera ocellata — described 1902: Biol. Cent. Amer. Coleop- 
tera, v. 4, pt. 4, p. 3, would appear from the figure and descrip- 
tion to be related to Phy. eximins, except that the beak is 
described as short and widened at the tip. The species has 
eleven black elytral spots and ochreous and gray scales. The 
two specimens were collected at "Omilterre, Guerrero, 8,000 
feet (H. H. Smith) Mex." 

The following names are undoubtedly nomina niida but if 
opportunity offers someone should examine the specimens 
provided they are still in existence and correctly determine 

1837: Dejean: Cat. Coleop. Coll. Dejean. p. 286. Phy- 
tonomus conjusus — Amer. boreal. 

1869: Giebel: Col. Univ. Halle Wittenberg, p. 44, No. 28, 
"Hypera nudirostris Germar in litt., Nord Amer." No. 31 an 
unknown species from "Nord Amer." 

Hypera punctata Fabricius. 

1762: Geoffroy: Ins. 1: 279. "Curculio no. 5." 

1779: Schaeffer: Icones Insectorum, tab. 25, fig. 6. 
Curculio punclalus: 

1775: Fabricius: Systema Entom., p. 1.50, no. 119. 

1781: Fabricius: Species Insectorum, 1: 190, no. 166. 

1781: L5iichart: Verz. u. Besch. d. Tyrol Ins. Kafer, 1 (pt 1): 16, no. 221 

1787: Fabricius: Mantissa Ins., 1: 117, no. 221. 

1787: Schneider: Neus Mag. Entom. 3(pt 2): 116, no. 190. 

17SS: Zschachii: Pars Entom. p. 21, no. 458. 

1789: Villers: Entom. Fauna Suec, 1:21,3, no. 169; 4: 279, no. 169. 

1790: Gmelin: Linn. Syst. Nat., ed. xiii, p. 1786, no. 378. 

1790: Olivier: Hist. Nat. Ins., 5: 541, no. 315. 

1790: Rossi: Fauna Etrusca, 1: 131, no. 335. 

1792: Fabricius: Entom. Syst. emend. l(pt 2): 472, no. 329. 

1792: PaykuU: Monog. Cure. Suec, p. Ill, no. cvii. 

1795: Fabricius: Nomen. Entom., p. U. 

1795: Herbst: Nat. Ins. Kafer, 6:,505, no. 540. 

1795: Panzer: Entom. Germanica, p. .329, no. 174. 

1795; Rossi: Fauna Etrusca, (Hellwig ed.) 1: 1,39, no. 335. 

1795: Weber: .Nomen. Entom. sec. E. S. Fab. p. 57. 

1796: Fabricius: Index Alphabeticus, E. .S. emend., p. 54. 

1797: Bergstrasser: Epit. Entom. Fab. Nomen. p. 68, 71. 

1800: Paykull: Fauna Suecica, 3: 306, no. cxxix. 

1801: Fabricius: Svs. Eleutherat., 1:529, no. 1.33. 

1805: Illiger: Magaz. f. Insektenkundc, 4: 1.33. 

1828; Boitard: Man. d'Entom., 1: 409. 

1853: Moretti: in Gene, ed. 2. 
RhvtirhatiHs punctalus: 

' 1802: Fabricius: Ind. Gen. et sp. Svs. Eleuth., p. 69, no. 54. 

1813: Gvllenhal: Ins. Suec, 1 (pt3): 108, no. 38. 

1820: Billberg, Enum. Ins., p. 42. 

1827; Gyllenhal: Ins. Suec, l(pt 4, app. 3): 572, no. 38. 
Brachvrhinus punctalus: 

1.804; Latreille: Hist. Nat. Gen. et pare. 11; 171, no. 59. 

1911] Hypera and Phytonomiis in America 397 

Hvpera punctata: 

1817: German Germ. &- Zincker Mag. 2: 340. 

1821: Germar: Germ. & Zincker Mag., 4: 344, no. 22. 

1821: Dejean: Cat. coll. Coleop., p. 89. 

1826: Curtis: Brit. Entom. 2: no. 116, 1 (fixes type of genus). 

1826: Sturm: Cat. Ins. Sammlung, 1: 157. 

1829: Curtis: Guide Arr. Brit. Ins., p. .50, no. 3. 

1829: Stephens: Sys. Cat. Brit. Ins., p. 167, no, 1707. 

1831: Stephens: Entom., 4; 93. 

1848: Walton, Ann. Mag. Nat. Hist. (2) 1: 297. 

1849: Walton: Stett. Ent. Zeit., 10: 259. 

1861: Waterhouse: Cat. Brit. Coleop., p. 71, no. 1. 

1863: Lacordairc: Hist. Nat. Ins. Coleop., 6: 401. 

1869: Giebel: Verz. z. Mus. Univ. Halle-Wittenberg p. 43, no. 1. 

1869: Targione-Tozzetti: Bui. Ent. See, Ital., 1: 80-81. 

1869: Kraatz: Verz. Kafer Deutsch., p. 52. 

1871: Kirsch: Berl. Ent. Zeit. 15: 184. 

1871: Gemmingcr & Harold: Cat. Coleop., 8: 2386. 

1877: Stein & Weise: Cat. Col. Europe, ed. 2, p. 143. 

1880: Koppen: Die Schadlichen Ins. Russlands, p. 209. 

1880: Rupertsberger: Biol, die Kafer Europa, p. 200. 

1883: Weise in H. R. &■ W. Cat. Col. Eur. et. Cau., ed. 4, p. 159. 

1884: Bargagli: Bui. Ent. Soc. Ital., 16: 170-1. 

: Rass. Biol. Rinc. Europei, p. 97. 

1888: Bedel: Col. Bassin de la Seine, p. 255. 

1889: Fauvel: Rev. Ent., 8: 157. no. 458. 

1890: : Bui. Ent. Soc. Ital., 22: 275. 

1891: Fowler: Brit. Coleop., 5: 229, 231. 

1891: Schneider: Coleop. & Lepidop. Bergen, p. 112, no. 34. 

1891: Weise in H. R. & W. Cat. Col. Eu. Cau. et Arm. Ross., p. 303. 

1893: Bertolini: Bui. Ent. Soc. Ital. 25: 244, no. 6. 

1894: V. d. Hoop: Tijd. v. Entom., .37: 172. 

: Eckstein: Deutsch Fisch. Zeit. Stettin., p. 

1903: Everts: Coleop. Nccrlandica, p. 600. 

1908: Torka: Entom. Blatter, 4: 77. 

1909: Cecconi: Rev. Col. Ital., 7: 46. 
Phylonomus punctatus: 

1826: Schonherr: Cure. Dispos. meth., pt. 4, p. 175. 

1829: Gebler: in Lededour Reise, p. 168. 

1830: Gebler: Bemerk uber d. Ins. Sib. vorz. Altai, p. 168. 

1833: Carmagnola: in Villa: Cat. Coleop. dupl. p. 24. 

1833: Dejean: Cat. Coleop. coll. Dejean, ed. 2. p. 264. 

1834: Schonherr: Gen. et sp. Cure. 2 (pt 2): 401. 

1837: Dejean: Cat. Coleop. coll. Dejean. ed. 3, p. 287. 

1839: Falderman: Neue Mem. Soc. Nat. Mosc, 6: 189. 

1842: Schonherr: Gen. et sp. cure, 6 (pt 2): 346. 

1843: Sturm: Cat. Kafer Sammlung, p. 201. 

1843: .Schmidt: Stett. Entom. Zeit., 4: 23. 

1844: (Dohm:) Cat. Coleop. Eur., p. 52. 

1847: Hoehhuth: Enum. Russelkafer, Kaukasus u. Transk., p. 491, no. 98. 

1849: Gaubil: Cat. Syn. Coleop. Eu. et Alg., p. 156, no. 3. 

1849: Redtenbachcr: Fauna Austriaca, Die Kafer, pp. 433-4. 

1849: (Dohm): Cat. Col. Eu., p. 61. 

1851: Ferris: Mem. Ac. Sc. Lyon, n. s. 1:373. 

1853: Murray: Cat. Col. Scotland. 

1853: Zebe: Syn. der bisher Deutsch. aufgef. Coleop. p. 75. 

1855: Jac. du Val.: Gen. Coleop. d' Europe, p. 109. 

1857: Lentz: Neue Verz. der Preuss. Kafer, p. 124. 

1858: Dohrn: Cat. Col. Eur., p. 78. 

1858: Mathieu; Ann. Ent. Soc. Belg., 2: 197, no. 188. 

1858: Redtenbachcr: Fauna Austriaca, Die Kafer, ed. 2, p. 727. 

398 Annals Entomological Society of America [Vol. IV, 

1862: Schaum: Cat. Col. Europa, ed. 2, p. 89. 

1864: Jekel: Ann. Ent. Soc. Fr. (4) 4: 562, 563. 

1865: Disconzi: Entom. Vicentia, p. 79, 81, no. 36. 

1865: Thomson: Skand. Coleop., 7: 101. 

1866: de Marseul: Cat. Coleop. Eur. ct. conf., p. 100, no. 2. 

1867: Capiomont: Rev. d. Hyperides, p. 121-3, pi. 11, fig. 12; 12, f. 1. 

1868: Capiomont: Rev. d. Hyperides, p. 421. (sep. p. 201). 

1868: Villa: Rel. sugli Ins. d. Trifogli. 

1868: Villa: La Lombardia, 13 giugno. 

1874: Siebke: Enumeratio Ins. Norvegicum, fasc. 1, p. 264-5. 

1877: Hevden: Jahrb. Nassau. Vereins, 29: 311. 

1878: Sclineider & Leder: Beit, kennt. Kauk. Kaferfauna, p. 287. 

1879: Targione-Tozzetti: Ann. Agr. Minis. Agr. & Com. p. 30. 

1881: Heyden: Cat. Col. Sibiria, p. 165. 

1881: Riley: Amer. Naturalist, 15: 750-1. 

1881: Lintner: Husbandman (Elmira, N. Y.), 14 Sep., p. 3, 6, 7. 

1881: Lintner: Cultivator and Coun. Gent., 46: 031, 29 Sep. 

1881: Lintner: Cultivator and Coun. Gent., 46: 647, 6 Oct. 

1881: Riley: Amer. Naturalist, 15: 912-4. 

1882: Leconte: Tr. Am. Ent. Soc. 9: proc. p. xxxvi. 

1882: Riley: Amer. .\aturalist, 16: 248-9. 

1882: Riley: 12th Rep. Ent. Soc. Ont., p. 17. 

1882: Lintner: 41st Rep. N. Y. St. Agr. Soc., pp. 40-50. 

1882: Lintner: Rural New Yorker, 41: 010, 9 Sep. 

1882: Rilev: Report of the Entomologist, p. 111-9, pi. 10, fig. 1. 

1883: Riley: in Rep. U. S. Dept. Agr. f. 1881-2, pp. 171-9, pi. 10, f. 1. 

1883: Lintner: First Rep. St. Entom. N. Y., pp. 247-53, fig. 

1883: Lintner: Rural New Yorker, 42:310, 19 May. 

1883: Mann: Psyche, 4: (Bibliog. rec. 3309, 3325, 3326). p. 96. 

1884: Lintner: Countrv Gentleman, 49: 457, 29 May. 

1884: Lintner: Ont. Co. Times (N. Y.) 29 May, 4 June. 

1884: Devereaux, Clvde Times (N. Y.) 29 May. 

1884: Lintner: Tr. N. Y. St. Ag. Soc. f. 1877, v. 33:221-34, fig. 

1884: Kilman: Canad. Entomologist, 16: pp. 144-5. 

1884: Lintner: Canad. Entom., 16: 182. 

1884: Moffat: Canad. Entom., 16: 215. 

1884: Sanders: Can. Entom., 16:209-10. 

1885: Henshaw: List Col. of Amer. N. of Mex., p. 137, no. 8227. 

1885: Lintner: 2nd Rep. St. Entom. N. Y., pp. 3, 14-15. 

1885: Kilman: 15th Ann. Rep. Ent. Soc. Ont., p. 32. 

1885: Sanders: loth Ann. Rep. Ent. Soc. Ont., p. 152. 

1886: Arthur: Botanical Gazette, 11: 14-17, pi. 2 (Enlomophlhora). 

1886: Arthur: 4th Ann. Rep. N. Y. Agr. Exp. Sta., f. 1885, p. 241-65, fig. 10. 

1886: Rilev: Proc. Ent. Soc. Wash., 1: 20. 

1888: Forbes: Psyche, 5: 10, 45-6. 

1888: Lintner: 4th Rep. St. Ent. N. Y., p. 178. 

1888: Thaxter: Mem. Bost. Soc. Nat. Hist. 4: 172-175, pL 17, figs. 200-21. 

1889: Kilman: Canad. Entom.. 21: 136. 

1889: Henshaw: Bibliog. Ec. Entom. pt. iii. 

1889: Hamilton: Tr. Am. Ent. Soc. 16: 155, no. 454. 

1889: Lintner: 5th Rep. St. Ent. N. Y., p. 272, fig. 40. 

1889: Lintner: 4th Rep. N. Y. St. Museum, pp. 151, 244. 

1889: Smith: 10th Rep. N. J. Agr. E.xp. Sta., p. 241, 313. 

1890: Liebeck: Entom. News, 1: 12. 

1890: Rilev & Howard: Ins. Life, 3: 70-71. 

1890: Smith: Cat. Ins. New Jersey, p. 2.50. 

1890: Lintner: 6th Rep. St., Entom. N. Y., p. 182. 

1891: Smith: Ins. Life, 3: 231-2. 

1891: Lintner: 7th Rep. St. Entom. N. Y. p. 315, fig. 33. 

1891: Smith: Ins. Life, 4: 270. 

1891: Hamilton: Ins. Life, 4: 270. 

1911] Ilypcra and Phylononius in America 399 

1891: Osborn: Orange Judd Farmer, 10 Oct.. p. 229. 

1891: Seidlitz: Fauna Transsvlv., p. 674. 

1891: Weed: Ins. & Insecticides, pp. 235-7, tig. 126. 

1892: Webster, F. M.; Ohio Farmer, 4 June, p. 422. 

1892: Riley & Howard: Ins. Life, 4: 40i. 

1892: Rilev & Howard: Ins. Life, 5: 54. 

1892; Smith: Ins. Life, 5:98, 

1892: Webster: Ins. Life, 5: 99. 

1893: Butler: Indiana Farmer, 14 Jan., figures. 

1893: Riley &: Howard: Ins. Life, 5: 279. 

1893: Lintner: Country Gentleman, 58: 380, 18 May. 

1893: Webster, F. M.: Ins. Life, 6: 186. 

1803: Lintner: Sth Rep. St. Entom. N. Y., p. 300. 

1893: Lintner: 9th Rep. St. Entom., N. Y., p. 440. 

1893: Riley & Howard: Ins. Life, 6:328. 

1893: Webster: Canad. Entom. 25: 255. 

1893: Meinert: Fort. z. Mus. Billc Larver Coleop., p. 279, no. 508. 

1894: Riley & Howard: Ins. Life, 6: 328. 

1894: Webster, F. M.: Ohio Farmer, 10 May, p. 377, figs., 17 May, p. 39. 

1S94: Wheeler: Bui. 110, Mich. Agr. Exp. Sla., p» 47-52, fig. 1-3. 

1894: Hopkins: Ins. Life, 7: 145. 

1894: : Michigan Farmer, 8 Sep. 

1894: Webster, F. M.: Ins. Life, 7: 203. 

1894: Hr)ward: Insect Life, 7: 273. 

1894: Buckhout: Rpt. Pa. Agr. Exp. Sta. f. 1893, p. 92. 

1895: Lintner: 10th Rpt. St. Ent. N. Y., pp. 448, 511. 

1895: Schwarz: Bui. 6, Div. Ornith. & Mam., U. S. Dept. Agric, p. 62. 

1895: Henshaw: Bibliog. Am. Ec. Ent., pt 4, auth. A-K inc. 

1895: Webster: Ohio Farmer, 23 May, p. 417; .30 May, p. 437, fig.; 13 Je, p.__477. 

1896: Henshaw: Bibliog. Am. Ec. Ent., pt 5. auth. L-Z inc. 

1896: Webster: Bui. 68, Ohio Ag. Exp. Sta.. pp. 27-31, fig. 

1896: Kellicott: loum. Columbus (O.) Hort. Soc. pp. 48^9. 

1896: Smith: Economic Entomology, pp. 229-30, figs. 234, 235. 

1897: Johnson: Bui. 9, n. s. Div. Ent. U. S. Dept. Agr., pp. 80-82. 

1897: Packard: Psyche, 8: 125. 

1897: Webster: fourn. N. Y. Ent. Soc, 5:202. 

1898: Banks: Bibliog. Am. Ec, Ent., pt 6, 1888-1896 inc. 

1898: Beutenmuller: Journ, N. Y. Ent. Soc, 0:40. 

1898: Webster: (Jhio Farmer, 19 May, p. 403. 

1898: Felt: Country Gentleman, 26 May, 63: 406. 

1898: Howard: Yearbook f. 1897, U. S. Dept. Agr. p. 529. 

1898: Johnson: Bui, .57, Md. Agr. Exp. Sta.. p. 110. 

1898: lohnson: Bui. 17, n. s. Div. Ent. U. S. Dept. Agr., pp. 92-94. 

1898: Felt: Rpt. St. Ent. N. Y., pp. 246, 257, .329, 336, 337, 374, 378. 

1899: Hunter: .Ann. Rpt. Neb. St. Bd. Agr. p. 24. 

1899: Kilman: 20th Ann. Rpt, Ent. Soc. Ont., pp. 90-91. 

1899: Sanderson: .National Rural, 2 Nov. 

1899: Lugger: 5th Rpt. St. Ent. Minn., p. 189-90, figs. 198, 199. 

1899: Lugger: Bui. 66, Minn. Agr. Exp. Sta., p. 85. 

1900: Smith: Cat. Ins. New Jersey, p. 343, figs. 154, 155. 

1900: Chittenden: Bui. 22, n. s. Div. Ent. U. S. Dept. Agr., p. 57. 

1900: Johnson: Bui. 20, n. s. Div. Ent. U. S. Dept. Agr., p. 96. 

1900: Fletcher: Bui. 26, n. s. Div. Ent. U. S. Dept. Agr. p. 96. 

1901: Evans: 31st Rpt. Ent. Soc. Ont., p. 34. 

1901: Fletcher: 31st Rpt. Ent. Soc. Ont., p. 62. 

1901: Fletcher: Rpt. Ent. &• Bot. Dom. Canada, p. 

1901: Banks: Bibliog. Ec. Ent., pt 7, 1897-1899 inc. 

1901: Sanderson: 28th Rpt Del. Agr. Exp. Sta. p. — . 

1901: Petri: Monogr. Coleop. Trib. Hyperini, pp. 120, 202, pi. 3, fig. ID. 

1901: Petri: Bestim. Tab. Coleop. Hft 44, Hyperini, pp. 19, 39. 

1902: Felt: 17th Rpt. St. Ent. N. Y., p. 845. 

400 Annals Entomological Society of America [Vol. IV, 

1002: Osborn, Bui. 37, Div. Ent. U. S. Dept. Agr., pp. 115, 116. 

1902: Sanderson: Ins. Inj. Staple Crops, pp. 177-79, fig. 98. 

1902: Ulke: Proc. U. S. Nat. Mus., 25: 33, 55 (Dist. Col. list). 

1902: Webster: 32nd Rpt. Ent. Soc. Ont., p. 64. 

1902: Durv: Joum. Cin. Soc. Nat. Hist., 20: 107, 182 (sep. p. 1) 

1902: : Nuove Relaz. Staz. Ent. Agrar. Firenze. 

1903: Chittenden: Yearbook f. 1902, U. S. Dept. Agr. p. 728. 

1903: Balkwill: 33rd Rpt. Ent. Soc. Ont., p. 44. 

1903: Bethune: 3.3rd Rpt. Ent. Soc. Ont., p. 44, 98. 

1903: Fletcher: 33rd Rpt. Ent. Soc. Ont., p. 44, 98. 

1903: Felt: 18th Rpt. St. Ent. N. Y., p. 123. 

1903: Sempers: Inj. Insects (W. A. Burpee Co.) 17th ed., p. 178, fig. 155. 

1903: Tower: Zool. Jahrbuch. 7 (pt 3): 520. 

1904: Titus and Pratt: Bui. 47, Bur. Ent. U. S. Dept. Agr., p. 67. 

1904: Felt: 19th Rpt. St., Ent. N. Y., p. 184. 

1905: Banks: Bibliog. Am. Ec. Ent., pt. 8, 1900-1904, inc. 

1905: Currie: Bui. ,53, Bur. Ent. U. S. Dept. Agr., p. 32. 

1906: Fletcher: 36th Rpt. Ent. Soc. Ont., p. 84. 

1906: Folsom: Entomology, ed. 1, pp. .58, 381. 

1906: Weise in H. R. & W.: Cat. Coleop. Eur., ed. 2, p. 655. 

1907: Close: 16-18th Rpts Del. Agr. Exp. Sta., p. 106. 

1907: Pierce: Ann. Rpt. Neb. St. Bd. Agr., p. 258. 

1907: Webster: Bui. 69, Bur. Ent. U. S. Dept. Agr., pp. 55, 89. 

1908: Pierce: Bui. 79, Bur. Ent. U. S. Dept. Agr., p. 41. 

1908: Houghton: Joum. Ec. Ent., 1: 297, 298, 299. 

1908: Fletcher: 3Sth Rpt. Ent. Soc. Ont., p. 127. 

1908: Sanderson: Ins. Inj. Staple Crops, ed. — , pp. 177-9, fig. 98. 

1909: Webster, R. L.: Entom. News, 20: 81. 

1909: Folsom: Bui. 134, 111. Agr. Exp. vSta., pp. 15.5-164, figs. 13-16. 

1909: Bur. Ent.: Yearbook f. 1908, U. S. Dept. Agr., p. 509. 

1910: Webster. R. L.: Joum. Ec. Ent., 3: 502. 

1910: Pettit: Bui. 258, Mich. Agr. Exp. Sta., pp. 44-46, figs. 11, 12. 

1910: Titus: Bui. 110, Utah Agr. Exp. Sta., p. 19. 

1910: Titus: Joum. Ec. Ent., 3: 460. 

1910: Smith: Cat. Ins. N. Jersey, p. 381, fig. 157. 

1911: Webster, F. M., in Westgate and Hillman, Farmer's Bull. 455, p. 39, 
fig. 22. 
jOurculio austriacus: 

1781: Schrank: Enumer. Insect. .A.ustriae, p. 123, no. 334. 

1784: Herbst: Fues. Archiv., 5: 82, no. 70. 

1789: Villers: Entom. Suec. Fauna., 1: 214, no. 177; 4: 281, no. 177. 

1790: Gmelin: Linn. Svst. Nat. ed. xiii, p. 1778, no. 324. 

1790: Olivier: Hist. Nat. Ins. 5: 571, no. 56. 

1795: Herbst: Nat. Ins. Kafer, 6: 243, no. 207, T. 77, fig. 11. 

1795: Panzer: Fauna Ins., civ. no. 8. 

1800: Paykull: Fauna Suecia Insecta, 3: .306, no. cxxix. 

1802: Marsham: Entom. Brit., 1: 302, no. 184. 
Brachvrkinus austriacus: 

1804: Latreille: Hist. Nat. Gen. et pare, 11: 179, no. 89. 
R}iync}ia:nus austriacus: 

1813: Panzer: Ind. Fauna Germ., p. 200, no. 21. 

1813: Gyllenhal: Ins. Suecica, 1 (pt 3): 108, no. 38. 

1819: Samouelle: Entom. Useful Comp., p. 369. 

1819; Samouelle: Nomen. Brit. Ent. Alphab. Arrang., 1: 36. 

1820: Billberg: Enum. Insect., p. 42. 
Hypera punctata var. austriaca: 

1871: Gemminger & Harold: Cat. Coleop., 8: 2386. 
Phvlonomns punctatus var. austriacus: 

' 1901: Petri: Monogr. Coleop. Trib. H^rperini, p. 202. 

1901: Petri. Bestim.-Tab. Coleop. Hft. 44, Hyperini, p. 39. 

1911] Hypera and Phytonomus in America 401 

Curculio piclus: 

1785: Fourcroy: Entom. Paris., 1: 117, no. 5. 

1789: Villers: Entom., Fauna Suec, 1; 216, no. 187; 4: 282, no. 187. 
Hypera puiiclala var. picta. 

1871: Gemminger & Harold: Cat. Coleop., 8: 2386. 
Curculio linzensis: 

1790: Gmelin: Linn. Syst. Nat. ed. xiii, p. 1799, no. 477. 
Hypera punctala var. linzensis: